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HARVARD UNIVERSITY 



Ernst Mayr Library 

of the Museum of 

Comparative Zoology 




Volume 9 • 2001 



. 



MCZ 
LIBRARY 

JUL. 2 jj 2002 

HARVARD 
UNIVERSITY 



I i mi Zhao 
Chengdu Institute of Biology. Academia Sinica, Chengdu. Sichuan. China 



Associate Editors 



Ki ll.ii Autumn 
Lewis & Clark College, Portland. Oregon. USA 



Theodore J. Papenfuss 

Museum of Vertebrate Zoology. University of California. 
Berkeley. California. USA 
J. Robert Macey James Ford Parham 

Department of Biology. Washington University. St. Louis. Museum of Paleontology. University of California. 



Missouri. USA 



Berkeley. California. USA 



Editorial Board 



Kraig Adler 

Cornell University. Ithaca. New York. USA 

Natalia B. Ananjeva 

Zoological Institute, Si. Petersburg. Russia 

Steven C. Anderson 

University of the Pacific. Stockton. California. USA 

Aaron Bauer 

Villanova University. Villanova. Pennsylvania. USA 

Christopher Bell 

University of Texas. Austin. Texas. USA 

Leo Borkin 

Zoological Institute, St. Petersburg. Russia 

Bihui Chen 

Anhui Normal University, Wuhu. Anhui. China 

I .|iiiiiii Cheng 



Xiang Ji 

Hangzhou Normal College. Hangzhou. Zhejiang, China 

Pi-peng Li 

Yantai Normal College, Yantai. Shandong. China 

Ronald Marlow 

University of Nevada. Las Vegas. Nevada. USA 

Robert W. Murphy 

Royal ( )ntario Museum. Toronto. Ontario. Canada 

Giiren NiLson 

University ol Goteborg. Goteborg. Sweden 

Nikolai Orlov 

Zoological Institute, St Petersburg, Russia 

Hidetoshi Ota 

Department of Biology, University of the Ryukyus. Nishihara, 

Okinawa. Japan 

Soheila Shafii 



Institute of Marine Biology. National Taiwan Ocean University. University of Shahid Bahonar. Kerman. Iran 

Keelung, Taiwan. China 

Ilya Darevsky 

Zoological Institute. St. Petersburg. Russia 

Indraneil Das 

Madras Crocodile Bank, Vadanemmeli Perur. Madras. India 

William K. Duellman 

University of Kansas. Lawrence. Kansas. USA 



Hajime Fukada 

Sennyuji Sannaicho. Higashiyamaku. Kyoto. Japan 

Carl Gans 

University of Michigan. Ann Arbor. Michigan. USA 

Robert F. Inger 

Field Museum. Chicago, Illinois, USA 



Hai-tao Shi 

Hainan Normal University. Haikou. Hainan. China 

\iu-ling Wang 

Xinjiang Normal University. Urumqi. Xinjiang. China 

Yue-zhao Wang 

Chengdu Institute of Biology. Academia Sinica. Chengdu. 

Sichuan. China 

Yehudah Werner 

Hebrew University, Jerusalem. Israel 

Ken-tang Zhao 

Su/hou Railway Teacher's College, Suzhou. Jiangsu. China 



Asiatic Herpetological Research is published by the Asiatic Herpetological Research Society (AHRS) and the Chinese 
Society for the Study of Amphibians and Reptiles (CSSAR) at the Museum of Vertebrate Zoology. University of California. 
The editors encourage authors from all countries to submit articles concerning but not limited to Asian herpetology. All 
correspondence outside of China and requests for subscription should be sent to AHR. Museum of Vertebrate Zoology. 
University of California. Berkeley. California. USA 94720. or by email to asiaherp@uclink.berkeley.edu. All 
correspondence within China should be sent to Ermi Zhao. Editor. Chengdu Institute of Biology. P.O. Box 416, Chengdu. 
Sichuan Province. China. Authors should consult Guidelines for Manuscript Preparation and Submission at the end of this 
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$5 per issue for surface mail or $10 per issue for air mail. Make checks or money orders payable in US currency to AHRS. If 
you do not have access to US currency, please notify us. and we will make other arrangements. 

Asiatic Herpetological Research Volume 9 succeeds Volume 8 (published in 1999). Vol. 7 (1997). Vol. 6 (1995). Vol. 5 (1993). Vol. 4 
( 1992). Vol. 3 (1990). and Chinese Herpetological Research Vol 2, which was published at the Museum of Vertebrate Zoology. 1988- 1 989 
as the journal for the Chinese Society for the Study of Amphibians and Reptiles. Volume 2 succeeded Chinese Herpetological Research 
1987. published for the Chengdu Institute of Biology by the Chongqing Branch Scientific and Technological Literature Press. Chongqing, 
Sichuan. China Acta Herpetologica Sinica ceased publication in June. 1988. 

Cover: Turtles of Turkey. Upper left: Trionyx triunguis. Dalyan. 6/16/00. Upper right: Tesnuio graeca anamurensis. Anamurium. Anamur, 
6/1 1/00. Lower left: Mauremys caspica rivulata Mamure Kalesi. Anamur. 6/10/00. Emys orbicularis. Lower right: Mamure Kalesi. Anamur. 
6/10/00. All photographs by Tonya Van Leuvan-Smith. 



2001 



Asiatic Herpetological Research 



Vol.9, pp. 1-5 



The Reproductive Biology of Rana boulengeri 

Wen-Jian Li 

Laboratory of Economic Frogs, Changde Teachers College, Changde, Hunan, 415000 China 

Abstract.- Maturation of Rana boulengeri Giinther occurs at the age of 24 to 36 months. After reproduction the 
gonads are in an intermittent period. Gonads begin to develop rapidly from next February to April. After May, the 
ovocytes grow mature in batches. In the wild, the adult frogs often spawn under waterfalls or in shallow pools 
where the water flows slowly, with high dissolved oxygen. The reproductive period lasts from May to August. 
The ovulation time of females and fertilization time are generally from 5:00 a.m. to 8:00 a.m. The Water 
temperature for deposition at the beginning is 15.5°C and the appropriate temperature for most deposition is 
17.7-22.5°C. The total duration under artificial constant temperature of 23±0.5°C lasts 261 hours. The 
appropriate temperature for development is 22-24°C. This species can not develop below 4°C, and it dies at 30°C. 
The development of the ovary is divided into 6 stages and the development of the testis is divided into 5 stages. 
The morphological and histological studies at different stages of sex cells are described. 

Key words.- Amphibia, Rana boulengeri, reproductive biology, China 



Introduction 

Rana boulengeri is a large frog that is found in the 
Wuling Mountains of Hunan Province. It is an impor- 
tant species for maintaining the ecological balance of 
the region (Yuan and Wen, 1990). Besides, it can be 
used as a kind of medicine (Li et al., 1993). In order to 
help protect this natural resource, these studies on its 
reproductive biology have been carried out. 

Material and Methods 

Laboratory experiment and artificial breeding 

The studies on the natural condition of habitat, growth 
and ages, characteristics of reproduction, the develop- 
ment of genital gland and development of the embryo 
of Rana boulengeri was conducted for four years 
from 1989 to 1993. 

Field investigation 

The field sites chosen for investigation were places 
where Rana boulengeri are common. Field observa- 
tions were made two or three times a month. 

Results and Discussion 

Habitat 

Natural habitat of Rana boulengeri. The Xiang Xi 
mountains are a part of the Wuling Mountains, which 
are located on the border of Hunan, Hubei, Sichuan 
and Guizhou; between north Latitude 27°44'- 29°48' 
and east longitude 109°10'-1 H"20'; with a mean ele- 
vation of 686 meters (the highest, 1900 meters above 



sea level; the lowest, 75 meters above sea level). The 
mountains are covered with evergreen broad-leaf sub- 
tropical forest. The mean air temperatures for a whole 
year are 16.10 degree-days. The temperature summa- 
tion above 10°C is 50320; the mean sun hours per 
year are 1292. 7h; frostless period is 270.6 days; the 
mean rainfall per year is 1397.2mm; the mean humid- 
ity per year is above 82% (R.H). These conditions 
constitute an ideal environment for Rana boulengeri. 

Rana boulengeri frequently live in brooks or 
ponds in mountain forests where there are few people. 
The water in the brooks or ponds is very clear. During 
the day, R. boulengeri often hide at the edge of small 
caves and during the night, they go out to feed. From 
July to September of 1987, the physical factors in 35 
observation points were measured. Their values are: 
the elevation of the habitat is 204-675 meters above 
sea level; air temperature 17.2-24°C; water tempera- 
ture 14.9-20.5°C; the depth of water 0.2-1.0 meters; 
the transparency of water 0.2-1.0 m; the humidity 
92.0-97.5%. 

Activities of Rana boulengeri and temperatures of 
habitat. Because the air temperature and water tem- 
perature play important roles in the development and 
growth of R. boulengeri; we observed their relation- 
ship in a cave (450m above sea level). In autumn, 
when the water temperature fall lower than 12.5°C, R. 
boulengeri begins hibernation. In spring, when water 
temperature is above 12.5°C, they start to feed, and 
when water temperature is higher than 15.5°C, they 
enter the reproductive period. The air temperatures 
and water temperatures in all months are listed in 
Table 1. 



Vol. 9, p. 2 



Asiatic Herpetological Research 



2001 



Table 1 . Air and water temperatures of the habitat of 
Plana boulengen (450m above sea level). 



Month 



Mean air 
temperature 



Mean water 
temperature 



1 


8.1 


11.0 


2 


10.0 


12.2 


3 


10.8 


12.8 


4 


13.0 


14.4 


5 


14.6 


14.9 


6 


16.3 


16.1 


7 


23.9 


16.5 


8 


22.1 


16.3 


9 


18.1 


16.1 


10 


16.1 


12.9 


11 


13.1 


11.5 


12 


8.2 


10.0 



Chemical properties of the water. 

The chemical characteristics of the water inhabited by 
R. boulengen were measured and analyzed. The 
results are listed in Table 2. On the whole, the water is 
clear, thin, neutral and acidulous, with some contents 
of Ca, Mg and other minerals. 

Vertical distribution. Rana boulengeri are distrib- 
uted vertically from 200 to 700 meters above sea 
level, but most of them are found from 450 to 650 
meters. Older, large adult frogs are mostly found 
above 600 meters, while young frogs and tadpoles are 
found lower. 

Growth and age 

When bred in artificial pools, the mean body length 
and the mean body weight of the frogs just complet- 
ing metamorphosis are 19.8 mm and 0.95 g respec- 



tively. When feeding lasts to the end of the year, their 
body weight increases to 3.07 g and their body length 
to 31.2 mm. After feeding for 2 years, their body 
weight grows to 20.50 g, and the body length to 58.1 
mm. The growth curves of body weight and body 
length are logistic (i.e. "S" shaped) (Li, et al., 1993). 
In the field, it was found that the largest male was 261 
g with a body length of 134 mm. The largest female 
was 202 g with body length of 120 mm. 

Reproductive characteristics 

Environmental conditions of breeding areas. In the 

wild, the adult frogs often spawn under waterfalls or 
in shallow pools. The mean water area and mean 

water depth of natural spawn areas are 2.16 m" and 
0.35 m respectively. The water flows slowly with high 
dissolved oxygen. The mean value of pH is 6.3. Gen- 
erally, there are small stones, sand, ratty plant and 
leaves, or humus on the bottom. There are bryophytes 
and algae on the bottom as well as on the sides of the 
pools. 

Reproductive period. In west Hunan at 563 m above 
sea level, R boulengeri begin reproduction in May and 
end in August. The reproductive peak is in June and 
July. Female frogs may have three clutches, the egg 
number of the second and third clutches depend on 
the availability of food and environmental conditions. 

Temperature. When water temperature reaches 
15.5°C R. boulengeri begins to spawn. The optimum 
temperatures for large numbers of females to spawn 
are 17.7-22.5°C. From July to August, the mean air 
temperature of natural spawning sites is 21.4°C, and 
the optimum humidity is above 95%. 

Reproductive behavior. In middle or late March, R. 
boulengeri end their hibernation and feed for a month. 
Then the male and female frogs aggregate at spawn- 
ing fields. Before mating, male frogs call for about 15 
days as a courting period. The ovulation time of 
females and fertilization times are generally from 
5:00 a.m. to 8:00 a.m. When the female spawns, she 
creeps slowly and her cloaca is near the walls of pool 
or brooks. 



Table 2. Analysis of the water inhabited by R. bou/engeri(mglL). Date of analysis: 5, January, 1990; elevation where 
water was sampled: 204m above sea level. 



PH 


total hardness 


Ca 


Mg 


Cu 


Zn 


Fe 


Hg 


Mn 


Cd 


6.71 


12.92»C 


58.36 


13.66 


0.013 


0.183 


0.793 


0.001 


0.036 


0.003 


Pb 


Ag 


Co 


Cr 


Do 


Sulfate 


N(N0 2 ) 


N(HN0 3 ) 


N(NH 4 ) 


Water type 


0.0148 


0.002 


0.017 


0.0199 


4.45 


9.89 


0.001 


0.026 


0.075 


Thin acidulous 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 3 



> 
o 

•B 7 



9? 
o 

e 

a> 
o 
o 

<U 

o5 



3- 




J I L 



J I I I l_L 



10 11 12 1 2 3 4 5 6 7 8 9 month 

Figure 1 . The seasonal change of Mature coefficient 
of ovary (1991-1993) 

Eggs. The mean number of eggs spawned in each 
batch is about 218. The egg of R. boulengeri is big 
with a mean diameter of 3.98 mm, enveloped by 3 
layers of gum membrane (The diameter of egg includ- 
ing membrane is 15.85 mm). The outer layer is the 
thickest, the inner layer the thinnest. The outer layer is 
very sticky, so that many eggs connect together to 
form a long cluster, the end of egg cluster is stuck on 
the wall or to ratty plants under water. 

Development of genital glands 

Development of ovary. The development of ovary 
can be divided into 6 stages. 

Stage 1. Multiplication period of ovogonium: The 
body weights (BW) of young frogs are 7.3-12.5 g and 
the body lengths (BL) are 37.2-43.6 mm. The ovogo- 
nium cells, aggregating densely with the ability of 
division, have not been surrounded by follicle cells. 
Stage 2. Growth of ovocyte (oocyte): When the BW 
of young frog are 10.2-19.5 g and BL are 47.2-56.2 
mm, ovogonium cells develop into ovocyte cells. The 
ovocyte clusters are divided by connective tissue of 
the ovary and surrounded by follicle cells. 

Stage 3. Transition growth of ovocyte: When BW of 
young frog are 36.5-74.2 g and BL are 71.7-84.3 mm, 
the volume of ovocyte cells increases. The follicle 
cells increase from 1 to 2 layers and the zona pellu- 
cida becomes clear. 

Early stage 4. Formation of yolk in ovocyte: When 
young frogs weigh 58.7-80.8 g and their BL are 81.5- 
89.7 mm, oocytes begin rapid growth and particles of 
yolk begin to form. 



Late stage 4. Ovocyte cells filled with yolk: When 
frogs have BW of 79.7-125.8 g and BL of 87.3-98.0 
mm, the ovum cells are fully filled with yolk and the 
nuclei are isolated in the middle of the cells. The two 
layers of follicle cells and zona pellucida are well 
marked. 

Stage 5. Slanting of nucleus of oocyte: When BW are 
86.3-161.5 g and BL are 89.4-1 12.3 mm, the nucleus 
inclines to the side of animal pole. At this stage, the 
female can ovulate under the action of gonadotrophic 
hormone. 

Stage 6. Ovary after ovulation: When frogs body 
weights decrease to 70.6-102.5 g and body length 
decrease to 82.1-100.2 mm. In this stage, corpora 
lutea appear in ovary. 

Development of testis. The development of testis can 
be divided into 5 stages. 

Stage 1. Multiplication period of spermatogonium: At 
this stage, spermatogonium aggregates densely, and 
ranges irregularly. 

Stage 2. Formation of sperm-tube: In this stage, the 
spermatogonium is separated into sperm-tube by con- 
nective tissue of testis and part of spermatogonium in 
sperm-tube becomes spermatocyte of the first order. 
Stage 3. Differentiation of spermatocyte: In this stage, 
the number of spermatogonium in sperm-tube 
decreases. On the wall of sperm-tube, there are pri- 
mary sperm mother cells, secondary sperm mother 
cells and spermatophore formed by spermoblast 
respectively. 

Stage 4. Formation of sperm: This is the ripe stage of 
the testis. The spermatogonium and the spermatocyte 
of the first order are few on the wall of sperm-tube 
instead of spermatocyte of the secondary order and 
spermatophore formed by spermoblast respectively. 
After the forming of spermatozoon, the spermatozoon 
aggregates densely first, then disperses. 

Stage 5. Post-ejecting: Most spermatozoons have 
been ejected and only a few could be found in the 
sperm-tube. 

Female order of ovary development and age at sex- 
ual maturity. After metamorphosis, the ovaries of the 
young frogs reach stage 1 in 2-3 months; stage 2 in 6- 
8 months; stage 3 in 10-13 months; early stage 4 in 
21-24 months; late stage 4 and stage 5 in 33-36 
months (if ecological conditions are very suitable, in 
only 22-24 months), when the females can spawn nat- 
urally or artificially. After spawning, the ovaries 
reaches stage 6. 

Male order of testis development and age at sexual 



Vol. 9, p. 4 



Asiatic Herpetological Research 



2001 



to 

& 



1.2 



1.1 



1.0 



0.9 



0.8 



ti 0.7 



c 

CD 

'o 

0) 
o 
o 

a) 

g 

ra 



0.6- 



0.5 




A 



J I I I L 



I I . 



10 11 12 1 2 3 4 5 6 7 8 9 month 

Figure 2. The seasonal change of Mature coefficient 
of testis (1991-1993) 

maturity. After metamorphosis, the testes of males 
develop into stage 1 in 2-4 months; stage 2 in 8-10 
months; stage 3 in 14-18 months; stage 4 in 24-26 
months. At this stage the males can mate with the 
females. 

Type of spawning. In the sexually mature females, 3- 
4 grades of oocyte can be seen. This is the cytological 
proof of multi-oviposition of Rana boulengeri. 

Seasonal change of ovaries. The period from May to 
August is the reproductive time of R. boulengeri. 
Then the ovaries develop slowly and the mature coef- 
ficient of ovary (MCO) decreases to 3.10-3.35%. 
From February to April of the second year, the ovaries 
develop fast and the MCO reaches its highest value 
(5.73-14.5%). The mean value of MCO in May is 
8.47% (Fig.l). After May, the ovocytes mature in 
batches. 

Seasonal change of testes. The testes change little 
during the different seasons. The range of change of 
mature coefficient of testis (MCT) in a year is 0.55- 
1.14%. From April to July, testes develop faster than 
in other months. In July the MCT is 1.14%. In Octo- 
ber the MCT decreases to 0.55% (Fig.2). 

Embryonic development 

Li et al. (1993) reported in detail on the embryonic 
development of R. boulengeri. Under temperature 
conditions of 23±0.5°C, 261.05 ± 0.54 hours were 
required for the embryo to complete its development. 



The whole development procedure is divided into 25 
stags on the basis of both the morphological and the 
physiological changes during this period (Pollister 
and Moore, 1937; Shumway, 1940). The temperature 
summation above 0°C for the embryo to complete the 
whole development is 6004.15 degree-hours. 

Experiments have shown that temperature can 
affect the developmental rate of the embryo of R. bou- 
lengeri. Under conditions of 13°C, 16°C, 22°C, 24°C, 
26°C and 28°C, the time for the embryo to complete 
development is 937.97h, 687.78h, 321.61h, 304.76h, 
300. 18h and 295. 17h. In 2°C and 4°C water the 
embryo can not develop. In 7°C water the embryo 
develops very slowly. In 30°C water, the embryo can 
not complete development. The optimum tempera- 
tures for embryonic development are 22°-24°C (Li et 
al., 1994). 

Acknowledgments 

My research has been supported by the Asiatic Herpe- 
tological Research Society. I would like to thank Pro- 
fessor Er-mi Zhao (Chengdu Institute of Biology) for 
guiding my research and revising my paper. The 
author wishes to thank Changde Teachers' College for 
its financial support. 

Literature Cited 

Li, Hu-Ming, Wen-Jian Li, Xing-Guo Gong, and 
Ming-You Ma. 1993. Studies on the nutritional com- 
positions and energy content of the flesh of Rana bou- 
lengeri. Zoological Research, Kunming, 14(1 ):96. 

Li, Wen-Jian, Hu-Ming Li, and Ming Li. 1993. Pre- 
liminary studies on the artificial breeding of the young 
frog of Rana boulengeri Giinther. Pp. 255-259. In 
Zhao, Chen and Papenfuss (eds.), Proceedings of the 
First Asian Herpetological Meeting. China Forestry 
Press, Beijing. 

Li, Wen-Jian and Ming Li. 1993. The early embryonic 
development and stages of Rana boulengeri. Pp. 124- 
133. In Zhao, Chen, and Papenfuss (eds.). Proceeding 
of the First Asian Herpetological Meeting. China For- 
estry Press, Beijing. 

Li, Wen-Jian and Hu-Ming Li. 1994. Effect of temper- 
ature on the early embryonic development of Rana 
boulengeri. Pp. 508-512. In Proceedings of the sixti- 
eth anniversary of the founding of China Zoological 
Society. China Science and Technology Press, 
Beijing. 

Pollister, A. W and J. A. Moore. 1937. Tables for the 
normal development of Rana sylvatica.. The Anatomi- 
cal Record 68:489-496. 



2001 Asiatic Herpetological Research Vol. 9, p. 5 



Shumway W. 1940. Stages in the normal development 
of Rana pipiens. The Anatomieal Reeord 78: 1 39- 1 47. 

Yuan. Feng-Xia and Xiao-Bo Wen. 1990. A prelimi- 
nary study on living and feeding habits of Rana bou- 
lengeri in western Hubei Province. Chinese Journal of 
Zoology. Beijing 25(2)17-21. 



2001 



Asiatic Herpetological Research 



Vol. 9. pp. 6-8 



A New Species of Batrachuperus from Northwestern China 

Mingtao Song 2 , Xiaomao Zeng 1 , Guanfu Wu 1 , Zhijun Liu 'and Jinzhong Fu 3 

Chengdu Institute of Biology, Chinese Academy of Sciences. Chengdu. China 610041. Northern west Institute 

of Endangered Animals. Xi'an. China 710032. ~ Department of Zoology. University of Guelph. Guelph. Ontario. 

Canada NIG 2W1 

Abstract.- We describe a new species of salamander in the genus Batrachuperus from Tsinling Mts. in western 
China. The new species is morphologically most similar to B. longdongensis. but differentiated by the absence of 
horny covers on palms and tarsa. It represents the most northeastern distribution and the lowest elevation of the 
genus. 

Key words.- Caudata. Batrachuperus. new species. China. Tsinling 



Introduction 

The salamander genus Batrachuperus Boulenger. 
1878 contains seven species (Frost. 1985). Three spe- 
cies occur on the western side of the Tibet Plateau 
(Iran and Afghanistan). The remaining four species 
are found on the eastern side of the Tibet Plateau in 
China. During the tieldwork in 1986 and 1999. we 
collected specimens of Batrachuperus from Zhouzhi. 
China. This collection marks the most northeastern 
distribution of the genus. The specimens are different 



from all other described species (Fei et al., 1990). and 
we describe them as a new species. 

Batrachuperus taibaiensis new spe- 
cies (Fig. 1). Taibai Stream Sala- 
mander 

Holotype: NIEA 860122. An adult male from the 
upper stream of Heihe River, near Hua Er Ping Vil- 
lage. Zhouzhi County. Shaanxi Province, China 
(33.85°N, 107.82°E). collected by M. Song on August 
8. 1986. elevation 1260m. The holotype is deposited 




Figure 1 . Paratype of Batrachuperus taibaiensis (MVZ 230964). 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 7 



at the herpetological collection of Northwest Institute 
of Endangered Animals, Xi'an, China. 

Paratypes: Allotype. NIEA 860116, an adult 
female, collected from the same locality at the same 
time as the holotype. Other paratypes include NIEA 
860114-5, 860117-9, 860121, 860126-7, 860129-139 
and MVZ 230964-65, 230979-86. The NIEA series 
are collected from the same locality at the same time 
as the holotype and are deposited at the herpetology 
collection of Northwest Institute of Endangered Ani- 
mals, Xi'an, China. The MVZ series are collected 
from the same locality on September 30. 1999 by Z. 
Liu, and are deposited at the herpetology collection of 
the Museum of Vertebrate Zoology, University of Cal- 
ifornia. Berkeley. 

Diagnosis: A relatively large, robust stream sala- 
mander; distinguished from other members of the 
genus by the large size, vomerine teeth arranged in a 
"A" shape, and lack of horny cover on palms and 
tarsa. Morphologically, the species most closely 
resembles B. longdongensis. Both species are distin- 
guished from others by large size and vomerine teeth 
arranged in a "A" shape. Between the two species, B. 
longdongensis differs from B. taibaiensis by the pres- 
ence of the horny cover on palms, tarsa, and tail tip, as 
well as the presence of gill slits in adults. Batrachupe- 



rus taibaiensis is distinguished from geographically 
neighboring species, B. tibetanus, by its large size and 
the arrangement of vomerine teeth. Most phylogeneti- 
cally closely related to B. karlschmidti and B. tibeta- 
nus. Its cytochrome b gene sequence differs from B. 
longdongensis by 9.2%, from B. karlschmidti by 7%, 
and from B. tibetanus by 8.2-8.6% (J. Fu et al., 
unpublished data). 

Description: Batrachuperus taibaiensis is a large 
(adults males over 217 mm maximum SL), stout spe- 
cies. Head moderately depressed, its length from 
snout to gular fold longer than its width; snout short 
and round. Labial fold well developed, often partially 
covers the lower jar. Angle of jaw just behind the pos- 
terior corner of eye. Both maxilla and mandible with 
tiny teeth. Tongue elliptical, without free end. Ver- 
mine teeth four, arranged in "I \" shape. 

Body stout. Male body length slightly longer than 
tail length and female body and tail length about the 
same. Costal grooves 11. Limbs relatively short but 
strong; when adpressed, tips of digits do not overlap 
and are always separated by 1-3 costal folds in adults, 
but contacted in juveniles. Separation is greater in 
males. Fingers four, 3-2-4-1 in order of length. Toes 
four, 3-2-4-1 in order of length. Most individuals 
without palmar and tarsal tubercles. No horny cover 



Table 1 . Measurement (range, means) and percentage ratios of each character dimension to SVL. All measure- 
ments are in mm. 



Measurement 



Snout-vent length 
Tail length 
Head length 
Head width 
Forelimb length 
Hindlimb length 
Limb interval 
Vomerine teeth 



Holotype 



Allotype Male (n =11) 



Juvenile Female (n=9) 



102 



115 



26 



23 



26 



30 



52 



105 



105 



27 



25 



22 



30 



51 



92.9±12.1 

73—110 

98.5±13.7 

76—115 

25.4±2.9 

20—29 

20.4±1.6 

18—23 

22.7±2.5 

20—27 

29.6±3.0 

25—36 

44.8±7.9 

33—59 

4 



65 



60 



19 



13 



18 



21 



30 



102.1 ±7.1 

94—111 

101.3±7.5 

91—113 

27.2±1.5 

25—28 

21.4±1.7 

19—25 

23.7±1.2 

22—26 

30.9±2.5 

23—34 

52.2±8.7 

44—68 

4 



Vol. 9, p. 8 



Asiatic Herpetological Research 



2001 



on palms, tarsa, and ventral side of the fingers, and 
toes. Some individuals have horny cover of the very 
tips of the fingers and toes. Tail round at the base and 
gradually flattened laterally. Tail fin moderately high. 
Skin smooth. The measurements of the type speci- 
mens are presented in table 1. 

Habitat and distribution: This species is only 
known from the type locality, which is the most north- 
eastern distribution of the genus and the only known 
location from the northern slope the Tsinling Mts. It 
has also the lowest elevation of the genus (1260m). 

This species inhabits in fast moving streams (close 
to the headwater of Heihe River). During daytime, 
they are found under rocks in the stream, and have 
never been observed under rocks on the riverbank. 
The stream is well covered by the canopy from both 
sides of the river and has steep slopes. 

Etymology: The name taibaiensis is derived from 
the name of the nearby peak, Taibai Peak, which is the 
highest point of Tsinling Mts. 

Remarks: Recent phylogenetic study of the genus 
supported the species status of B. taibaiensis (Fu et al. 
unpublished data). It is the sister group of the clade 
including B. karlschimdti and B. tibetanus. Together, 



the clade is the sister group of B. yenyuansis and B. 
pinchonii. 

Acknowledgments 

We are grateful to E. Zhao, Z. Kou, Q. Wang and K. 
Li for their help. This research was supported by the 
Chengdu Diao Science Fund, Southwest Base Fund 
and NSFC 30070090 to X. Zeng and National Geo- 
graphic Society grant 6591-99 to J. Fu. 

Literature Cited 

Boulenger, G. A. 1878. Description de deux genres 
nouveaux de la famille des salamandrides. Bulletin de 
la Societe Zoologique de France 1878:71-72. 

Fei, L., C. Ye, and Y. Huang. 1990. Key to Chinese 
Amphibia. Chongqing Branch, Science and Technol- 
ogy Literature Publishing House, Chongqing, 
Sichuan. 

Frost, D. R. 1985. Amphibian Species of the World. 
The Association of Systematics Collections and Allen 
Press, Lawrence, Kansas. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 9-22 



A New Snake of the Genus Ho/ogerrhum Gur\\her (Reptilia; Squamata; 
Colubridae) from Panay Island, Philippines 

Rafe M. Brown 1 - 2 , Alan E. Leviton 3 , John W. Ferner 2 - 4 , and Rogelio V. Sison 5 

Section of Integrative Biology (C0930) and Texas Memorial Museum, University of Texas, Austin, Texas, 78712, 

USA. e-mail: rafe@mail.utexas.edu; Geier Collections and Research Center, Museum of Natural History and 

Science, 1301 Western Avenue, Cincinnati, Ohio 45203; - Department of Herpetology, California Academy of 

Sciences, San Francisco, California 94118, USA. e-mail: aleviton@CalAcademy.org; Department of Biology, 

Thomas More College, Crestview Hills, Kentucky, 41017, USA. e-mail: femerj@thomasmore.edu; Herpetology 
Section, Zoology Division, National Museum of the Philippines, Executive House, P. Burgos Street, Manila, 

Philippines, e-mail: nmuseum@webquest.com. 

Abstract.- We describe a new species of snake in the genus Hologerrhum from two forested localities in Antique 
Province, Panay Island, Philippines. To clarify species boundaries, we also redescribe its only known congener, 
H. philippinum, on the basis of historical collections and newly-acquired material from the Islands of Luzon, 
Marinduque, Polillo, and Catanduanes. The new species is the first Hologerrhum from the Visayan Aggregate 
Island Complex and differs from Hologerrhum philippinum in color pattern and scalation of head and body. The 
new species is one of several recently described vertebrates from Panay Island. Together, they indicate that 
forested regions of the individual islands of the Visayan Aggregate Island Complex (Negros, Panay, Cebu. 
Masbate, and other associated smaller islands) contain low levels of taxonomic endemicity that warrant further 
study. 



Key words.- Colubridae, Hologerrhum, Philippines, Panay Island, Visayas. 



Introduction 

Giinther (1858) erected the monotypic genus Holo- 
gerrhum to accommodate a single specimen from 
Hugh Cuming's Philippine collections that had been 
deposited in the Natural History Museum, London 
(Giinther, 1873; Fig. 1A). Giinther distinguished the 
new genus and species from members of the Philip- 
pine genus Cyclocorus by the presence of slight 
grooves in the enlarged fang-like teeth at the posterior 
end of the maxilla. Other slight differences between 
Hologerrhum and some species of Cyclocorus, not 
emphasized by Giinther but mentioned by other work- 
ers (Taylor, 1922a, 1922b; Leviton, 1965). include 
color pattern, slight scale pattern differences, and less 
strongly enlarged middle series of maxillary teeth in 
Hologerrhum. 

At the time of the description of Hologerrhum, no 
specific (island) locality data were available, but 
Giinther later (1879) referred a specimen from Placer, 
northeast Mindanao Island to this species. That speci- 
men (not seen by us) apparently is a representative of 
the genus Cyclocorus Taylor 1922c (vide Boulenger, 
1896; see also comment by Taylor, 1 922a: 1 16). 

Jan and Sordelli (1870) described Cyclochorus 
maculatus (generic name misspelled), on the basis of 



a specimen reportedly from Java Island, Indonesia 
(Fig. IB). Fischer (1885) followed by recognizing 
Cyclochorus lineatus var. maculatus reportedly from 
S. Mindanao Island, Philippines, but without refer- 
ence to new material. The type specimen of Cyclocho- 
rus maculatus later was shown to be a representative 
of H. philippinum (Giinther, 1873. 1879; Boettger, 
1886; Taylor, 1922a), suggesting locality errors by 
both Jan and Sordelli and Fisher. Later, Leviton 
(1965) inadvertently included Cyclochorus maculatus 
in the synonymy of Cyclochorus lineatus. 

Castro de Elera (1895) reported Hologerrhum 
philippinum from Baco, Mindoro Island. This impor- 
tant specimen could not be examined as it was 
destroyed during dissections by a biology class at the 
University of Santo Thomas, Manila (R. I. Crombie, 
personal communication) but the "Mindoro" locality 
information suggests a misidentification of a speci- 
men of C. lineatus (Taylor, 1922a; Leviton. 1963, 
1965). Griffin (1910) did not include Hologerrhum in 
his list of snakes from Polillo but did include the spe- 
cies in his key to the Philippine snakes (Griffin, 191 1), 
although he erred in attributing the type description to 
Boulenger and supplied no precise locality data. 



Vol. 9, p. 10 



Asiatic Herpetological Research 



2001 



It was not until E. H. Taylor's extensive work in 
the Philippines that specimens of H. philippimtm with 
reliable locality data became available. Taylor (1922a, 
1922b, 1922c, 1922d) consistently reported that this 
species was collected in montane habitats in primary 
forest and usually was associated with rocky stream 
beds on Luzon and its land-bridge satellite island of 
Polillo (Fig. 2). Still, Taylor (1922b:200) considered 
the species rare and only obtained eight specimens in 
two years of nearly continuous field work. 

During the nearly 80 years that have elapsed since 
Taylor's work, several additional specimens of H. 
philippinum have been collected on Luzon and its 
associated land-bridge islands (Marinduque, Polillo, 
and Catanduanes; see Specimens Examined; Fig. 2). 
During the same period, none have been found on 
Mindanao, Mindoro, or any of the other Philippine 
islands, thus bolstering the notion that H. philippinum 



is endemic to the Luzon Pleistocene Aggregate Island 
Complex (Fig. 2; sensu Heaney, 1986; Alcala, 1986; 
see also comments by Leviton, 1963). 

In 1992, while participating in the National 
Museum of the Philippines/Cincinnati Natural His- 
tory Museum Philippine Biodiversity Inventory 
(PNM/CMNH PBI), one of us (RMB) collected speci- 
mens of what appeared to be a distinctive new species 
of snake, similar to but obviously specifically distinct 
from H. philippinum, at 1025 m elevation on the west 
face of Mt. Madja-as, Panay Island. In addition to rep- 
resenting a previously unrecognized species, this 
specimen appears to be the first vouchered record for 
the genus on the Visayan Aggregate Island Complex 
(Fig. 2; Heaney, 1986; Hall, 1996, 1998). During the 
course of this study we examined all available US and 
Philippine museum collections of//, philippinum and 







■ ;;*•, 







Figure I.The first illustrations of Hologerrhum. (A) Gunther's (1879) drawings of H. philippinum 'and (B) Sordelli's 
plate of Cyclochorus maculatus (= H. philippinum) from Jan and Sordelli (1870). 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 



Less than 1 20 m 
submarine contour 



200 



km 



Luzon - 



Mindoro- 
Busuanga jj 




* Balanes islands 



Babuyan islands 



Polillo 

Mannduque 
..■■' 

Catanduanes 



Palawan 

.10" 



Leyle 



Mindanao 



• Hologerrhum philippinum if Hologerrhum dermali 

Figure 2. Map of the Philippine islands with the major 
Pleistocene aggregate island platforms indicated by 
the position of the 120 m underwater bathymetric con- 
tour (following Heaney, 1986); known collection locali- 
ties of the two species of Hologerrhum are indicated. 
Darkened circles represent collection localities for 
Hologerrhum philippinum 'Gunther, 1858, stars are 
localities for Hologerrhum derma/mew species. 

Hologerrhum n. sp. In this paper, we redescrihe H. 
philippinum and describe the new species from Panay. 

Material and Methods 

We surveyed the forested slopes of Mt. Madja-as from 
6 April- 1 May 1992, utilizing elevational transects as 
described by Ruedas et al. ( 1994) and as modified by 
Brown et al. (1995, 1996, in press; Ferner et al., this 
issue). Sampling techniques employed a variety of 
search and trapping methods, including multiple-per- 
son time-constrained searches, sticky traps, and pit- 
fall traps (Simmons. 1987). Specimens were fixed in 
I07t buffered formalin and transferred to 70% ethanol 
approximately 2 months following preservation. 

Museum specimens (see Specimens Examined, 
below) were examined for color pattern and scale 
character differentiation and only data scored by 
RMB and AEL were used in an effort to minimize 
inter-observer sources of variation (Lee. 1990). We 



used Dowling's (1951a. 1951b) methods for counting 
scales and expressing scale row reduction formulae, 
and applied the Evolutionary Species Concept (Simp- 
son. 1961: Wiley. 1978; Frost and Hillis. 1990) in 
making taxonomic decisions. 

Species accounts 

Hologerrhum philippinum Gunther 1858 
Figures 3-6 

Hologerrhum philippinum. Gunther ( 1 858: 1 86). 

Cyclochorus maculatus, Jan and Sordelli 
(1870:36; generic name misspelled; the specimen 
illustrated is H. philippinum with doubtful locality 
data). 

Hologerrhum philippinum. Gunther (1873: 171: 
specimen is a member of the genus Cyclocorus. vide 
Boulenger. 1896). 

Cyclochorus lineatus var. maculatus Fischer. 
1885:81. 

Hologerrhum philippinum, Boettger (1886:115) 
Castro De Elera (1895:438: specimen probably 
Cyclocorus lineatus): Boulenger (1896:33); Taylor 
(1922a:116; I922b:200, 1922c:138); Ross and 
Gonzales (1991:67); Brown et al. (1996:13). 

Hologerrum philippinum Griffin. 1911:263 
(generic name misspelled). 

Diagnosis: H. philippinum differs from its conge- 
ner, H. dermali, by (1) the presence of 12-30 (vs. 7- 
10) pairs of alternating black spots on nuchal region 
and anterior dorsum. (2) a pale orange to salmon (vs. 
bright yellow) venter, (3) absence (vs. presence) of a 
black midventral stripe. (4) labials cream or yellow 
(vs. labials bright white with thin midlabial black 
stripe). (5) dorsum tan to orangish brown or dark 
brick red (vs. dark purplish brown), (6) chin and 
throat of adults pale tan to orange, immaculate or with 
faint white spots in some specimens (vs. darker pur- 
plish brown with black and white ocelli), (7) invari- 
able presence of moderate to enlarged pretemporal 
(length more than half that of secondary temporal; vs. 
pretemporal reduced or absent), and (8) posterior tips 
of parietals extend caudally, posterior parietal suture 
forming a medially inverted V-shaped cleft (vs. poste- 
rior portions of parietals squared off. with no medial 
cleft). 

Description: Body cylindrical, ventrals convex, 
head slightly distinct from neck, not flattened; eyes 
small, pupil round: vertebral ridge absent. 

Rostral scale much broader than deep, scarcely 
visible from above, subtriangular with ventromedial 
groove in dorsal aspect: nasal divided, naris piercing 



Vol. 9, p. 12 



Asiatic Herpetological Research 



2001 




Figure 3. Live photograph of Hologerrhum ph///pp/numUom the Zambales Mountains of western Luzon Island 
(female, PNM 2490; photograph copyright D. Wechlser). 



A 



B 






-« . •A-*"*' 



I 



£5 



¥ 



D 



- ,>_ 



* 




$&■* 



Figure 4. (A) Dorsal and (B) lateral view of the head of Hologerrhum dermali : (CMNH 5075); (C) dorsal and (D) lat- 
eral view of the head of Hologerrhum phi/ipp/num (PUM 2490). 



2001 



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Vol. 9, p. 13 



Table 1 . Summary of diagnostic characters distinguishing Hologerrhum dermal! {new species; from Panay Island) 
from Hologerrhum philippinum (Gunther, 1858; from the Luzon Aggregate Island Complex). 



Characters 



H. philippinum 



H. dermali 



Pretemporals 

Ventral nuchal blotches 



enlarged 
-, + 



reduced or absent 



Lateral black ventral spots 


-, + 


+, fused into line 


Dorsal nuchal spots 


12-30 


7-10 


Midlabial stripe 


- 


+ 


Dorsal live color 


tan to reddish brown 


purplish brown 


Ventral live color 


orange to salmon 


bright yellow 


Adult throat color 


tan to orange (some spotted white) 


purplish brown w 
ocelli 


Midventral dark stripe 


- 


+ 


Parietal suture 


notched 


unnotched 


Ventral scale at 2nd dorsal 
reduction 


97-112* 


84-97 



Subcaudals (females) 



42-53 



57-61 



* Excluding apparently aberrant counts of CAS 61554 (Table 2). 



suture between pre- and postnasal; together, nasal 
scales square to triangular; dorsal border of nares 
formed by thick shelf of prenasal, ventral border 
formed by extension of postnasal; internasals as long 
as broad, slightly shorter than prefrontals, laterally 
contacting both pre-and postnasals, forming a vague 
right triangle with 45° face oriented anterolaterally; 
loreal single, as large as or only slightly smaller than 
ventral preocular, half as high as postnasal, pentago- 
nal, surrounded by postnasal, lateral edge of prefron- 
tal, dorsal and ventral preoculars, and second 
supralabial; prefrontals longer than internasals, with 
irregular lateral extensions caused by presence of con- 
cave curved suture with preoculars (concave surface 
oriented posterolaterally); frontal twice as long as 
broad, longer than to equal to its distance to the end of 
the snout, shorter than parietals; anterolateral corner 
of frontal barely contacting medial point of preocular 
(e.g., CAS 31553, 60951, MCZ R-25693-94, PNM 
6505), or with substantive contact between frontal and 
preocular squeezed off by contact between supraocu- 
lar and prefrontal (e.g., CAS 60950, 61554, 62430, 
134075; PNM 2120, 2490; USNM 498718, MCZ R- 
25695); posteromedial point of frontal extends past 
posterior margin of supraoculars for distance shorter 
than or equal to length of internasals; supraoculars 
very large, nearly as long as and slightly narrower 



than frontal; parietals very large, laterally contacting 
dorsal postocular, pretemporal, and highly enlarged 
secondary temporal, together bordered posteriorly by 
three (CAS 61554, 134075), four (CAS 60950, 
61553; USNM 498718, MCZ R-25695) or five (CAS 
60951, 62430; PNM 2490, MCZ R-25693-94) undif- 
ferentiated nuchals; posterior tips of parietals extend 
caudally, parietal suture forming a distinct medially 
inverted V-shaped cleft, in which a single slightly 
enlarged to undifferentiated nuchal (Fig. 6) lies; tem- 
porals arranged in three to four irregular vertical rows 
with enlarged posttemporals extending caudally 
beyond posterior ends of parietals; temporal formu- 
lae: (L) 2/1 + 1/1+2, (R) 1 + 1 + 1/1+2 (PNM 2490), (L, 
R) 1+2+1/1+2 (CAS 60950), (R, L) 1 + 1+2+3 (MCZ 
R-25695), (L) 1 + 1+2+3, (R) 1 + 1+2+2 (CAS 60951), 
(L) 1+1+2+3, (R) 1 + 1+2+4 (MCZ R-25694) (L) 
1 + 1 + 1/1+2, (R) 1 + 1+2+3 (CAS 61554, PNM 6505), 
(L, R) 1+1+1/1+2 (CAS 62430, 61558, 134075; 
USNM 318363, 498718; TNHC 60114, MCZ R- 
25693, PNM 2120), (L) 1 + 1+2+3, (R) 2+1/1+2 
(USNM 319037), (L); pretemporal relatively 
enlarged, its length much more than half that of sec- 
ondary temporal. 

Orbit surrounded by supraocular, two preoculars 
(dorsal larger than ventral), two postoculars, and 
supralabials 3-5; supralabials eight, fifth largest; 



Vol. 9, p. 14 



Asiatic Herpetological Research 



2001 



Table 2. Scale row reduction formulae (Dowling, 1951b) variation in H. philippinum philippinum (Gunther, 1858; 
from the Luzon Aggregate Island Complex) and the type series of H. derma// (new species; from Panay Island). 



Specimen (sex) 



Reduction 1 



Reduction 2 



Hologerrhum phillippinum 



3+4=3(4) 

CAS 6095 l(juv.) 19 17 

4+5=4(5) 



3+4=3(107) 

17 15 

3+4=3(106) 



4+5=4(5) 

CAS 60950 (juv.) 19 17 

3+4=3(4) 



3+4=3(103) 

17 15 

3+4=3(105) 



3+4=3(7) 

CAS 61553 (0 19 ' 17 

4+5=4(5) 



3+4=3(109) 

17 15 

3+4=3(112) 



3+4=3(7) 
CAS 61554(0 19 17 

4+5=4(6) 



3+4=3(93) 

17 15 

3+4=3(95) 

+4(96) 

3+4=3(100) 



3+4=3(7) 

CAS 62430 (juv.) 19 17 

3+4=3(6) 



3+4=3(101) 

17 15 

3+4=3(103) 



3+4=3(16) 

PNM 2490(0 19 17 

5+6=5(5) 



3+4=3(110) 

17 15 

4+5=4(109) 



5+6=5(5) 

PNM 2120(0 19 17 

5+6=5(6) 



3+4=3(95) 

17 15 

-4(99) 



3+4=3(4) 
PNM 2120 (juv) 19 n 

3+4=3(4) 

3+4=3(6) 

USNM 3 19037 (0 19 17 

3+4=3(5) 



3+4=3(97) 

17 15 

-4(96) 

3+4=3(105) 



17- 



3+4=3(109) 



-15 



3+4=3(5) 

USNM 318363(0 19 17 

3+4=3(7) 



3+4=3(100) 

17 15 

3+4=3(99) 



3+4=3(5) 

USNM 498718 (m) 19 17 

3+4=3(5) 



3+4=3(5) 
TNHC 60114(0 19 17 



3+4=3(8) 



-4(100) 

17 15 

^4(105) 

3+4=3(102) 



17- 



-4(100) 



-15 



4+5=4(4) 
MCZ R-25693 (0 19 17 

3+4=3(4) 



3+4=3(100) 

17 15 

3+4=3(100) 



2001 



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Vol. 9, p. 15 



Table 2. (continued) 



Specimen (sex) 



Reduction 1 



Reduction 2 



4+5=4(4) 

MCZR-25694(0 19 17 

4+5=4(4) 



3+4=3(102) 
17 15 

3+4=3(103) 



5+6=5(8) 

MCZ R-25695 (juv) 19 : 17 

5+6=5(6) 



3+4=3(97) 

17 15 

3+4=3(100) 



Mean 



x = 6.3 ± 3.2 SD; n = 12 



x =5.5±1.2SD;n=12 



x = 102.2 ± 4.9 SD; n = 12 
x =103.6 ±4.9SD;n=12 



Hologerrhum dermali 



3+4=3(5) 

PNM2711 H (f) 19 17 

3+4=3(6) 



3+4=3(94) 

17 15 

3+4=3(92) 



-4(3) 

PNM 6505 p (f) 19 17 

3+4=3(6) 



-4(97) 

17 15 

-4(92) 



3+4=3(5) 

CMNH 5075 p (f) 19 17 

3+4=3(7) 



3+4=3(84) 

17 15 

3+4=3(84) 



Mean 



x =4.3±1.2SD;n = 3 



x =6.3 ±0.6SD;n=12 



x =91.7±6.8SD;n=12 
x. =89.3±4.6SD;n = 12 



" = Holotype; p = Paratype 

infralabials eight; mental subtriangular, with highly 
pointed posterior tip caused by medially concave 
curved suture with first infralabial; first infralabials 
differentiated, elongate, with curved medial points 
nearly contacting anterior to genials; second infralabi- 
als reduced, squarish; infralabials 2-5 increasing dra- 
matically in size (fifth largest in verntral aspect), then 
decreasing sharply to infralabial 8; infralabials 1^ in 
contact with anterior genials, 4—5 in contact with pos- 



terior genials; mental groove distinct and broad; sub- 
labials thin, followed medially by 3^4 similarly-sized, 
longitudinal rows of gulars, medial two pairs (anterior 
to first ventral) slightly enlarged; number of gular 
pairs between posterior genials and first ventral two 
(CAS 61553) or three (remaining specimens). 

Dorsals smooth, without apical pits, vertebrals 
undifferentiated from paravertebrals, in 146-176 (X- 
157.4 ± 8.1 SD; n = 12) transverse rows on body, 42- 



Vol. 9, p. 16 



Asiatic Herpetological Research 



2001 





Figure 5. (A) Dorsal and (B) ventral view of a paratype of Hologerrhum dermalt '(CMNH 5075); (C) dorsal and (D) 
ventral view of Hologerrhum philippinum (PNM 2490). 



56 (JC = 49.3 ± 5.3 SD; n = 12) on tail; first scale row 
reduction (i.e., reduction of 19 to 17 scales around 
body; Table 2) occurring at point on body correspond- 
ing to ventrals 4-16 (left: X = 5.5 ± 1.2 SD; n = 12; 
right; X = 6.3 ± 3.2 SD; n = 12), second (17 to 15) 
occurring between ventrals 93-1 10 (left: X - 103.6 ± 

4.9 SD; n = 12; right X = 102.2 ± 4.9 SD; n = 12); 
ventrals broad, each slightly angulated laterally, 136— 
158 (X= 146.7± 6.4 SD; n = 12); subcaudals 42-56 

(x = 47.5 ± 4.6 SD; n = 12). The single adult male 
specimen (USNM 498718) has 156 vertebrals, 149 
ventrals, 56 caudals, and 55 subcaudals. Anal undi- 
vided; tail with enlarged vertebral row (dorsocaudals) 
formed by fusion of midvertebral row with both flank- 
ing paravertebral rows. Hemipenes of USNM 498718 
are extremely narrow and elongate, and are covered 
with uniformly minute spines; hemipenes extend in 
situ to the 14th subcaudal plate. 

Measurements (in mm): SVL 251-347 mm (X = 
280.8 ± 47.1 SD) for ten mature females; tail length 
X = 56-96 (X = 73.8 ± 15.0 SD) for eight mature 
females with complete tails. 

Coloration in preservative: Dorsum tan, orang- 
ish-tan to brown, with 12-30 (X = 21.8 ± 7.5 SD; n = 
12) alternating dark brown to black spots (Figs. 1A, 3, 
5C), each with three associated small white spots 
(Fig. 3), fading in intensity posteriorly, where they are 



replaced on scale rows 4-5 by a pair of dorsolateral 
black lines, gaining intensity posteriorly and continu- 
ing to tip of tail; faint vertebral thick gray stripe (1-3 
scales in width) becoming increasingly apparent pos- 
teriorly from midbody; a pair of light cream lines dor- 
sal (medial) to black lines; posterior (distal) portions 
of each dorsal scale slightly to markedly darker than 
remainder of scale; dorsal occiput colored as body 
(PNM 2490) or slightly darker (CAS 60950) to mark- 
edly darker (CAS 134075); melanic pigment congre- 
gated on medial suture between parietals, on posterior 
portion of frontal, and on lateral edge of head; distinct 
longitudinal dark midnuchal stripe evident from pos- 
terior edge of parietals to second pair of nuchal spots 
(Fig. 1A), occasionally (e.g., juveniles CAS 134075 
and MCZ R-25695, adult female USNM 318363) 
very dark and forming a distinct nuchal cross (Fig. 
IB); one specimen with a pair of bright white nuchal 
spots immediately anterior to nuchal cross (USNM 
319037; Fig. 1A); lateral aspects of head colored as 
dorsal, with distinct thin black line dorsally bordering 
supralabials (Fig. 4D) and stretching from tip of snout 
to just beyond supralabial 8; labials creamy yellow to 
tan, occasionally with a few black flecks (CAS 60950) 
or with ventral half of labials dark gray (USNM 
319037); venter immaculate cream to pale yellow or 
orange; each ventral with dark lateral pigment in the 
form of a small black spot or brown to black longitu- 
dinal bar (Figs. 1A, 3), becoming a confluent black 
ventrolateral stripe on posterior portions of body and 
tail; some specimens with subtriangular black mark- 



2001 



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Vol. 9. p. 17 




> \ — ^ ' 

/ \ / \ 

I \^ / / / 



Figure 6. Comparison of the posterior portions of the parietal suture in (A) Hologerrhum philippinum (PNM 2490) 
and (B) Hologerrhum dermah '(CMNH 5075). 



ings on anterior 20 ventrals (USNM 319037. 498718; 
TNHC 60114); underside of head lighter (PNM 
2490). to distinctly darker (USNM 319037) than 
remainder of venter, especially in juvenile specimens 
(MCZ R-25694-95; CAS 134075) where ventral head 
coloration resembles that of adult Hologerrhum n. sp. 
(see below); tongue black with pale gray tips of bifur- 
cated portions. 

Coloration in life: (Fig. 3) Dorsum described as 
"bright reddish to orange brown, darker anteriorly" or 
"reddish brown" (Taylor 1922b:200) or "grayish 
brown on neck, fading to orangish brown posteriorly" 
(Brown et al.. 1996: 13); labials dirty white to creamy 
yellow; venter "uniform pale, orangish tan" (Brown et 
al.. 1996:13) to "bright reddish salmon, lighter anteri- 
orly" or "uniform coral to red" (Taylor. 1922b:200); 
underside of head pale orange to "dusky with milk 
white spots" (Taylor, 1922b:200). 

Hologerrhum derma//, n. sp. 

Figures 4-6 

Holotype: PNM 271 1. an adult female, collected 
at 0900 hr on 9 April, 1 992 by Rafe M. Brown at 1 5 1 
m above sea level in the area known locally as "Hang- 
gud Tubig" ("Big Water"), on the western face of Mt. 
Madja-as. Barangay Alojipan. Municipality of Culasi. 
Antique Province, Panay Island. Philippines (11°23' 
N. 122 09'E). 

Paratypes: CMNH 5075. an adult female, same 
data as the holotype except collected at 1030 hr on 6 
April 1992 by Rogelio V. Sison; PNM 3704. an adult 
female, collected February-March 1994 by Rogelio V. 
Sison at 750 m above sea level on Mt. Ranges, Sitio 



Banagon. Barangay Aningalan. Municipality of San 
Remegio, Antique Province, Panay Island, Philip- 
pines. 

Etymology: The specific epithet is chosen to 
honor Ronald "Dermal" Crombie, in recognition of 
his numerous contributions to Philippine herpetology 
and in thanks for the guidance he has provided RMB 
and JWF during the past several years of our work 
with Philippine amphibians and reptiles. 

Diagnosis: Hologerrhum dermali can be readily 
distinguished from its congener. H. philippinum. by 
(1) the presence of 7-10 (vs. 12-30) pairs of dark 
spots in nuchal and dorsal regions. (2) a bright yellow 
(vs. pale orange to reddish salmon) venter, (3) pres- 
ence (vs. absence) of a black midventral stripe, (4) 
bright white labials with midlabial black stripe (vs. 
cream or yellow labials; midlabial stripe absent), (5) 
dorsum dark purplish brown (vs. tan to dark orangish 
brown or dark brick red). (6) chin and throat of adults 
dark purplish brown with black and while ocelli (vs. 
pale tan to orange, immaculate or with faint white 
spots in some specimens). (7) pretemporal absent or 
much reduced (length less than half that of secondary 
temporal: vs. pretemporal invariably present and 
enlarged), and (8) posterior portions of parietals 
squared oft. with no medial cleft at parietal suture (vs. 
posterior tips of parietals pointed, extending caudally 
to form a medial inverted V-shaped cleft). 

Description of the Holotype: An adult female. 
Body cylindrical, ventrals convex, head slightly dis- 
tinct from neck, not flattened; eyes small, pupil round; 
vertebral ridge nonevideni. 



Vol. 9, p. 18 



Asiatic Herpetological Research 



2001 




Figure 7. Habitat of Hologerrhum dermalisX the type locality (following heavy rain). 



Rostral much broader than deep, barely visible 
from above, subtriangular with ventromedial groove 
in dorsal aspect: nasal divided, naris piercing suture 
between pre- and postnasal: majority of dorsal border 
of nares formed by prenasal. majority of ventral 
formed by postnasal: internasals as long as broad, 
slightly shorter than prefrontals, laterally contacting 
both pre- and postnasals, forming a vague right trian- 
gle with 45° face oriented anterolaterally; loreal sin- 
gle, distinctly smaller than ventral preocular. half as 
high as postnasal, pentagonal, surrounded by postna- 
sal, lateral edge of prefrontal, dorsal and ventral pre- 
oculars and second supralabial: prefrontals longer 
than internasals. with irregular lateral extensions 
caused by presence of concave curved suture with 
preoculars (concave surface oriented posterolater- 
ally): frontal twice as long as broad, longer than its 
distance to the end of the snout, a little shorter than 
parietals: frontal-preocular contact squeezed off by 
substantive contact between posteriolateral corners of 
prefrontals and anteromedial corner of supraocular: 
posteromedial point of frontal extends past posterior 
margin of supraoculars for distance shorter than 
length of internasals; supraoculars very large, nearly 
as long as and narrower than frontal: parietals very 
large, laterally contacting dorsal postocular, pretem- 



poral. and highly enlarged secondary temporal, 
together bordered posteriorly by five undifferentiated 
nuchals; posterior ends of parietals squared off. with 
no medially inverted V-shaped cleft at parietal suture 
(Fig. 6); enlarged posttemporals extend posteriorly 
only slightly beyond caudal margin of parietals; tem- 
porals (L) 1 + 1+2+3. (R) 1 + 1/1+2 (pretemporal much 
reduced, its length much less than half that of suture 
between parietal secondary temporal). 

Orbit surrounded by supraocular, two preoculars 
(dorsal larger than ventral), two postoculars, and 
supralabials 3-5; supralabials eight, fifth largest; 
infralabials eight; mental subtriangular. with highly 
pointed posterior tip caused by medially concave 
curved suture with first infralabial; first infralabials 
differentiated, elongate, with curved medial points 
nearly contacting anterior to genials; second infralabi- 
als reduced, squarish; infralabials 2-5 increasing dra- 
matically in size (fifth largest in ventral aspect), then 
decreasing sharply to infralabial 8; infralabials 1—4 in 
contact with anterior genials, 4-5 in contact with pos- 
terior genials; mental groove distinct; sublabials thin, 
followed medially by four similarly-sized, longitudi- 
nal rows of gulars, medial two pairs (anterior to first 
ventral) enlarged; two pairs of gulars between poste- 
rior genials and first ventral. 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 19 



Dorsals smooth, without apical pits, vertebrals 
undifferentiated from paravertebrals, in 140 trans- 
verse vertebral rows on body, 64 on tail; scale row 
reduction from 19 to 17 in nuchal region and from 17 
to 15 posterior to midbody (Table 2); ventrals 143, 
broad, each slightly angulated laterally; subcaudals 
61; anal undivided; tail with enlarged vertebrals 
formed by fusion of midvertebral row with both flank- 
ing paravertebral rows; SVL 220 mm; tail length 68 
mm. 

Coloration in preservative: Dorsum dark pur- 
plish brown with 10 tightly paired black spots, 
decreasing in size posteriorly (Fig. 5A) on anterior 
one third of body; caudal third of body with a pair of 
dorsolateral black lines (on scale rows 4-5) gaining 
intensity posteriorly and continuing to tip of tail; ver- 
tebral stripe absent; a pair of light, bright yellowish 
lines dorsal (medial) to black lines, especially bright 
on tail; posterior (distal) portions of each dorsal scale 
markedly darker than remainder; dorsal occiput col- 
ored as body; melanic pigment congregated on medial 
suture between parietals, on posterior half of frontal, 
and on lateral edge of head; supralabials bright white, 
dorsal border composed of thin black stripe (Fig 4B), 
from tip of rostrum to beyond angle of jaw; white 
labial coloration continues in the form of a broad 
white stripe to point opposite fifth ventral; midlabial 
thin black stripe (Fig 4B) continues posteriorly as 
ventral border of the white stripe in nuchal region; 
distinct dark brown midnuchal stripe evident from 
posterior edge of parietals to first pair of nuchal spots, 
very dark and confluent with nuchal spots, forming a 
distinct nuchal cross; chin and throat purplish brown 
with white circular spots encircled in black (ocelli) 
much like juvenile coloration in H. philippinum speci- 
mens; venter pale yellow with midventral thin black 
stripe, becoming interrupted on caudal portions of 
body, nearly obliterated by vent and continuing again 
caudal to vent for five ventrals; each ventral with dark 
lateral pigment in the form of a small black spot (ante- 
riorly) or black longitudinal bar (caudally), becoming 
a confluent black ventrolateral stripe on caudal por- 
tions of body and tail; tongue black with bright white 
tips on forked portions. 

Coloration in life: Dorsum and ventral surfaces 
of head light purplish brown, light areas dorsal 
(medial) to dorsolateral caudal lines medium yellow; 
labials bright milky white; venter very bright yellow 
with distinct black midventral stripe. Iris dark brown 
to brick red. 

Variation: One paratype (female, CMNH 5075, 
SVL 268 mm; tail 91 mm) has seven pair of dark dor- 
sal spots, slightly lighter midcephalic coloration and 



less yellow above the dorsolateral caudal black lines. 
The midventral black stripe continues to the tenth 
subcaudal. This specimen lacks the small pretempo- 
rals found in the holotype; temporals (R, L) 1 + 1/1+2; 
ventrals 143, subcaudals 60, vertebrals 156, dorsocau- 
dals 64 . The other paratype (female, PNM 3704, SVL 
327 mm; tail 93 mm) has nine pairs of nuchal spots, a 
faint midlabial line, and lacks midventral stripes on 
the subcaudals (present on anterior 2/3 of body). PNM 
3704 has the following counts (R, L) 1 + 1+2+3, 
1 + 1 + 1/1+2; ventrals 149; subcaudals 58; vertebrals 
155; dorsocaudals 57. Scale row reduction formula 
presented in Table 2. 

Ecology and habitat: The type of habitat in 
which H. dermali (Fig. 7) was collected on Mt. 
Madja-as has been classified as the transition zone 
between mixed dipterocarp (submontane) and mossy 
(upper montane) forests (Whitmore, 1984; Ferner et 
al., 1997). The forest consisted of two strata (a canopy 
of 10 m, and a subcanopy of 3-4 m with emergent 
trees as high as 18 m); herb and shrub layer vegetation 
was also abundant. The forest near the collection site 
was mossy and contained high densities of epiphytic 
ferns and orchids. Topography was qualitatively char- 
acterized as steep, with numerous valleys bordered by 
sheer rock escarpments and forest-covered ridges. The 
holotype was collected in a sun spot in the early after- 
noon in a rocky stream bed (10 m wide) with a central 
4 m wide channel of rapidly running water. The speci- 
men was basking 1 .5 m from water on the top of a flat 
rock. The Mt. Madja-as paratype was collected in the 
mid-morning and was crawling through leaf litter on 
the forest floor (30 m from the same stream) when 
captured. Paratype PNM 3704, collected in San 
Remegio, was found on the floor of secondary forest 
near a small dry stream bed. The circumstances of 
collection are very similar to those reported for H. 
philippinum on Luzon (Taylor, 1922b; Brown et al., 
1996; A. Diesmos, personal communication). 

Discussion 

The endemic Philippine genus Hologerrhum is now 
known to contain two species distributed on the 
Luzon and Visayan aggregate island complexes (Fig. 
2). There are no known Hologerrhum from the Pala- 
wan, Mindoro, Mindanao, Sulu Archipelago, or the 
Batanes faunal subregions (Fig. 2). 

The absence of any clear close relatives of Holo- 
gerrhum (Leviton, 1963, 1965) among SE Asian colu- 
brids renders speculations regarding the genus' 
affinities somewhat moot. However, we note that both 
Hologerrhum and Cyclocorus share characteristics 



Vol. 9, p. 20 



Asiatic Herpetological Research 



2001 



unique among Asian snakes, most notably, an 
unusual, presumably derived pattern of reduction in 
caudodorsal scale rows. In all species of Cyclocorus 
and Hologerrhum, caudodorsal reduction takes place 
by fusion of vertebral and paravertebral scale rows, 
resulting in an odd-numbered series of longitudinal 
rows of caudodorsals rather than an even number 
(characteristic of all other SE Asian colubrine snake 
genera known to us). The systematic affinities of the 
genus Hologerrhum are in need of further study. 

The description of Hologerrhum dermali brings 
the number of new species of vertebrates recently 
described the by the PNM/CMNH PBI team in the 
coastal Madja-as mountain range to six (Sison et al., 
1995; Gonzales and Kennedy, 1990, 1996; Brown et 
al., 1997; Ferner et al., 1997; Brown et al., 1999). 
Other collections from Panay contain at least three 
probable undescribed species of frogs and many other 
species of amphibians and reptiles of uncertain taxo- 
nomic status (many of which are, doubtlessly, unde- 
scribed species; Ferner et al., this issue). Most of these 
species presumably are reliant on the closed-canopy 
rain forests of the western portions of Panay. Accord- 
ingly, most should be considered severely threatened 
by deforestation (see Ferner et al., 1997:fig. 2). 
Recent survey work in the northwestern portions of 
Panay indicates that Hologerrhum dermali occurs in 
forested habitats at lower elevations as well as the 
montane localities reported here (M. Gaulke, personal 
communication). Unfortunately, the low elevation for- 
ests of Panay Island have nearly all been removed by 
an aggressive timber industry in the central Visayas. 
We expect that numerous other undescribed popula- 
tions of amphibians and reptiles will be discovered in 
Panay and the remainder of the Visayas if biologists 
are permitted access to these forests in order to cata- 
log and describe Philippine biodiversity. 

Specimens Examined 

Hologerrhum philippinum: Philippines, Luzon 
Island. Zambales Province, Municipality of Masinloc, 
Barangay Coto, 4.3 km N, 0.5 km E of Mt. High Peak, 
elevation 1550 m (15° 31' N, 120° 07' E): PNM 2490; 
Bataan Prov., Mt. Mariveles: CAS 60950-51; Isabela 
Prov., Municipality of Palanan, Barangay Didian, 
Sitio Natapdukan, elevation 50 m: PNM 6505; Kal- 
inga Prov., Municipality of Balbalan, Barangay Bal- 
balan: CAS 61553-54, MCZ R-25694; Caminares Sur 
Prov., Municipality of Naga City, Mt. Isarog, eleva- 
tion 900 m: USNM 31863; Cagayan Prov., Municipal- 
ity of Baggao, Barrio Santa Margarita, elevation 150 
m: USNM 319037, 498718; CAS 134075; Quezon 
Prov., Municipality of Tayabas, Barangay Camaysa, 



Mt. Banahaw, 1 150 m above sea level: TNHC 601 14 
Camarines Norte Prov., Municipality of Ruis, Baran- 
gay San Lorenzo, Mt. Labo Range: PNM 2120; 
Mountain Prov., Mt. Polis: PNM 67; Laguna Prov., 
Mt. Makiling: MCZ R-25695; Polillo Island, Polillo 
Prov., near town of Polillo: CAS 62430, MCZ R- 
25693; Catafiduanes Island, Municipality of Gigmoto, 
Barangay Summit Bordan, elevation 200 m: USNM 
319037. 

Hologerrhum dermali: See Holotype and 
Paratypes sections for this species. 

Acknowledgments 

For logistical assistance in the Philippines, we thank 
the Department of the Environment and Natural 
Resources (DENR), A. Alcala (Silliman University), 
P. Gonzales and R. Caberoy (PNM), R. Kennedy 
(CMNH) and the provincial DENR authorities of 
Antique Province. The Protected Areas and Wildlife 
Bureau of the DENR facilitated collecting and export 
permits necessary for the field portions of this study. 

For the loans of specimens or assistance while vis- 
iting museum collections, we thank the following 
individuals and their respective institutions (museum 
acronyms follow Duellman et al., 1978 and Leviton et 
al., 1985): J. Vindum, R. Drewes and J. Slowinski 
(CAS), R. Crombie, K. de Queiroz, and G. Zug 
(USNM), R. Kennedy (CMNH), R. Caberoy (PNM), 
A. Diesmos (De La Salle University), and D. Canna- 
tella (TNHC). Financial support for RMB's travel to 
CAS while working on this project was provided by a 
C. Stearns Fellowship of the California Academy of 
Sciences. We owe particular thanks to R. Crombie and 
M. Gaulke for their help and assistance and to D. 
Wechlser for providing live photographs of H. philip- 
pinum. 

Support for field work (by RMB, JWF) was pro- 
vided in part by the Zoology and Botany Departments 
and the College of Arts and Sciences of Miami Uni- 
versity (Oxford, Ohio), the Society for the Study of 
Amphibians and Reptiles, The Explorer's Club, the 
Department of Biology of Thomas More College, and 
the Cincinnati Museum of Natural History. The PNM/ 
CMNH PBI was supported by a grant (to R. Kennedy 
and P. Gonzales) from the John D. and Catherine T. 
MacArthur Foundation and by the benefactors of Cin- 
cinnati Musuem of Natural History. We thank L. 
Bockstanz, T. LaDuc, A. Gluesenkamp, T Devitt, A. 
Diesmos, and D. Cannatella for comments on earlier 
drafts of this manuscript. The description of Hologer- 
rhum dermali constitutes contribution No. 24 to the 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 21 



results of the PNM/CMNH Philippine Biodiversity 
Inventory. 

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Vol. 9. pp. 23-24 



The First Record of Ptyas korros (Co\ubr\dae) from Bangladesh 

M. Farid Ahsan 1 and Shayla Parvin 

Department of Zoology, University of Chittagong, Chittagong 4331, Bangladesh, e-mail: mfahsan@ctgu.edu 

Abstract.- This is the first report of Ptyas korros from Bangladesh. The specimens reported here were collected 
from the Cittagong University campus between the years of 1985 and 1996. They match previously described 
specimens of Ptyas korros except in the number of loreal scales. Ptyas korros is known to occur in the Assam 
region of India and so its presence in adjacent regions of Bangladesh is expected, but unconfirmed until now. 

Key words.- Serpentes, Colubridae, Ptyas korros, Bangladesh, distribution, biogeography 



The number of species of snakes occurring in Bang- 
ladesh is still debatable because of the lack of confir- 
mation of some species. However, lists have been 
compiled based on a number of field studies con- 
ducted in different parts of the country. Sarker (1975) 
recorded 20 species, Montaquim (1979) reported 18 
species, Montaquim et al. (1980) recorded 28 species 
and Rashid (1982) recorded 22 species from Bang- 
ladesh. Whereas Khan (1982) estimated the snakes 
species of the country as 78 and he later reported 79 
species in 1987 and 81 species in 1992 (of which the 
occurrence of two are doubtful). Sarker and Sarker 
(1988) reported 88 species, although some are uncer- 
tain. This report established the presence of one spe- 
cies, Ptyas korros (Schlegel, 1837). 

During the identification of the snake species pre- 
served in the Departmental Museum of Zoology, Chit- 
tagong University (CU), four specimens (three 




5 cm 



Figure 1. Specimen of Ptyas korros from Chittagong 
University Campus. 



collected from CU Campus [CUC] between 1985 and 
1996, and one not known but possibly from CUC) of 
P. korros were found (Fig. 1). Until now, nobody has 
reported the occurrence of P. korros in Bangladesh, 
although Husain (1977) predicted it. Its known range 
extends from Assam (India) through Myanmar 
(Burma) to Indo-China, Thailand, throughout the Ten- 
asserim to the Malay Peninsula, Indonesia (Sumatra, 
Java, Borneo), Southern China including Hainan, Tai- 
wan and Hong Kong (cf. Giinther, 1864; Smith, 1943; 
Zhao & Adler 1993). The occurrence of P. korros in 
the CUC, Chittagong, extends its distribution to Bang- 
ladesh. It might also occur in nearby parts of the 
country such as Jamalpur, Greater Sylhet, Chittagong, 
Chittagong Hill Tracts (Rangamati, Khagracheri and 
Bandarbans Districts). 

The specimens from the CUC are olive brown 
above and yellowish white below. The scales on the 
posterior part of the dorsal side of the body are edged 
with a dark brown pattern (looking black) producing a 
regular network pattern. The outer margins of the ven- 
trals and caudals are also sometimes edged with dark 
brown (less dark than the dorsal color). The coloration 
of the CUC specimens resembles the P. korros speci- 
mens described by Smith (1943). Smith (1943) diag- 
noses P. korros from Ptyas muscosus by the number 
of dorsal scale rows (15 in P. korros vs. 16 or 17 in P. 
muscosus) and the number of ventral scales (160-187 
in P. korros vs. 190-213 in P. muscosus). The CUC 
specimens have 15 dorsal scale rows and 177-188 
ventral scales. They differ from P. korros specimens 
described by Giinther (1864) in the number of loreal 
scales. Giinther reported two loreals, but three of the 
four CUC specimens have only one loreal. 

Literature Cited 

Giinther. A. C. L. G. 1864. The reptiles of British 
India. Oxford & IBH Publishing Co., New Delhi. 452 
pp + XXVI plates. 



Vol. 9, p. 24 



Asiatic Herpetological Research 



2001 



Husain, K. Z. 1977. Bangladesher banayajonju sham- 
pad O tar sangrakshan (Wildlife resources of Bang- 
ladesh and their conservation). Bangla Academy 
Bijnan Patrika. 3(3): 1-1 1. (in Benglai). 

Khan, M. A. R. 1982. Wildlife of Bangladesh -a 
checklist. Dhaka University Press, Dhaka. 173 pp. 

Khan, M. A. R. 1987. Bangladesher banayaprani 
(Wildlife of Bangladesh) vol. 1. Bangla Academy, 
Dhaka. 169 pp. (in Benglai). 

Khan, M. A. R. 1992. Bangladesher shap (Snakes of 
Bangladesh). Bangla Academy, Dhaka. 227 pp. (in 
Benglai). 

Montaquim, M. A. 1979. Snakes of several districts of 
Bangladesh. Unpublished M. Sc. Thesis, University 
of Dhaka, Dhaka. 63 pp + v plates. 

Montaquim, M. A., Sarker, A. H.; Khan, M. A. R., and 
K. Z. Husain. 1980. List of the snakes of Bangladesh. 
Bangladesh Journal of Zoology. 8(2): 127-130. 



Rashid, S. M. A. 1982. On some snakes of Bang- 
ladesh with notes on their habit, habitat, status and 
distribution. Unpublished M. Sc. Thesis, University of 
Dhaka, Dhaka. 108 pp + xv plates. 

Sarker, A. H. 1975. Snakes of Bangladesh. Unpub- 
lished M. Sc. Thesis, University of Dhaka, Dhaka. 

Sarker, M. S. U. and Sarker, N. J. 1988. Wildlife of 
Bangladesh (a systematic list with status, distribution 
and habitat). The Rico Printers, Dhaka. 69 pp. 

Smith, M. A. 1943. The fauna of British India, includ- 
ing Ceylon and Burma, Reptilia and Amphibia, vol. 3 
Serpentes. Taylor and Francis, London. 583 pp. 

Zhao, E. and Adler, K. 1993. Herpetology of China. 
Society for the Study of Amphibians and Reptiles, 
New York. 522 pp + 48 plates. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 25-29 



Influence of Temperature on Burrow Use by the Monitor Lizard Varanus 
panoptes o\ the Coastal Dunes at Fog Bay, Northern Australia 

Sean J. Blamires 

Morning Bell Language School, 671-3 Jukdo 2 Dong, Pohang City, Kyongsangbuk Do 791-052, South Korea. 

email: s_blamires@hotmail.com 

Abstract.- An increase in the number of Varanus panoptes burrows appearing among the sand dunes at Fog Bay 
in northern Australia was noticed during the dry season (June to August). Entrances of marked burrows were 
smoothed, and their interiors investigated using a burrowscope, to determine the monthly number of foraging and 
retreat burrows appearing over a 12 month period. In the dry season, increased numbers of both types of burrows 
were found. A temperature data logger was used to record temperatures of the sand surface, and at a depth of 50 
cm. Burrow depths were measured as how far the burrowscope could be lowered into each burrow. There was a 
positive correlation between the number of retreat burrows and burrow depths. There was a negative correlation 
between the number and depth of retreat burrows and the minimum sand-surface temperature and the maximum 
sand temperature at 50 cm. The number of foraging burrows was independent of sand temperatures. The results 
indicate a likely thermoregulatory use of retreat burrows with more and deeper burrows prevalent when overnight 
surface and subterranean sand temperatures decrease. 

Key words.- Reptilia, Varanidae, Varanus panoptes, Australia, Northern Territory, fossorial, thermoregulation 



Introduction 

Burrows serve many ecologically important purposes 
to many animals (Hansell, 1993). Lizards expend less 
energy throughout the day if a large portion of time is 
spent in burrows (Bennett and Nagy, 1977). Monitor 
lizards (Varanus spp.) utilise burrows for a variety of 
purposes such as thermoregulation (Cowles and 
Bogert, 1944), reducing water loss (Green, 1972), 
finding prey (Pianka, 1969), and for oviposition and 
retreat (Cowles and Bogert, 1944; Auffenberg, 1983). 

In a recent study of the monitor lizard Varanus 
panoptes (Fig. 1 ) inhabiting the coastal dunes at Fog 
Bay, Northern Territory, Australia (12°42'S; 
130°20'E), I detected a seasonal change in the num- 
ber of burrows present. A greater number of burrows 
appeared among the dunes in June-August (dry sea- 
son) compared to the rest of the year. These burrows 
also seemed quite deep compared to those seen in the 
wet season. 

Use of these burrows for oviposition is unlikely 
since Varanus panoptes produces eggs during the wet 
season. Free access to other water sources might rule 
out conservation of water loss. Overnight tempera- 
tures in the area often drop below 15°C in the dry sea- 
son. A likely explanation may be that the monitors 
retreat into burrows at a depth where sand tempera- 
tures remain high to conserve overnight body temper- 




Figure 1. Varanus panoptes'm the coastal dunes at 
Fog Bay, Northern Australia. 

ature, as has been found for some other monitors 
(Cowles and Bogert, 1944; Auffenberg, 1983). The 
monitors at Fog Bay inhabit the dunes and forage on 
the beach and dunes (Blamires, 1999). Increased for- 
aging activities may be another explanation for the 
number of burrows seen in the dry season. The aim of 
this investigation was to determine whether changes 
in the number of burrows between the wet and dry 



Vol. 9, p. 26 



Asiatic Herpetological Research 



2001 



seasons is a result of foraging, thermoregulation, 
retreat or a combination of these, or other, influences. 

Material and Methods 

The area of the investigation is approximately 5 km of 
beach along the northern-most mainland beach at Fog 
Bay (12°42'S, 130°20'E), approximately 80 km from 
Darwin, Northern Territory, Australia. The area is pri- 
vately owned and closed to the public. The sandy 
dunes are immediately backed by grassland domi- 
nated by spinifex vegetation and dispersed Pandanus 
trees. Black soil plains, mangroves and salt flats back 
the grassland. The dunes of the southern-most 1 km 
stretch of beach are backed by monsoon forest. The 
entire 5 km of beach, and the grassland and monsoon 
forest backing the dunes, was walked twice monthly 
over a 12 month period. 

All burrows encountered were marked with a 
depth of surveyor's tape tied to nearby vegetation 
(Fig. 2). To estimate visitation rates the entrances 
were smoothed and checked, on revisiting, for varanid 
tracks leading into it. 

A small video-camera device, called a "burrow- 
scope" (Dyer and Hill, 1991; Dyer and Aldworth, 
1998) was used to investigate the inside of burrows. 
The model I used was a modified version of that of 
Dyer and Aldworth, (1998). A small black and white 
CCD, 38mm x 38mm, camera (Samsung MOD-BW 
204), now popular for home security systems (Capel, 
1993) encased in a 375 ml jar was used. Infrared light- 
ing was used to illuminate the burrow for the camera, 
provided by 10 high intensity light emitting diodes fit- 
ted on a piece of Vero board. A small black and white 
video monitor (260 mm; 5.8 kg) was used for view- 
ing, operated directly from a 12V battery. The wiring 
from the camera to the monitor was approximately 2 
m long and encased in a garden hose. The depth of 
each burrow was determined by measuring the depth 
of hose that fed into the burrow to reach its end. 

Burrows were assumed to be for foraging when 
were noted to intersect the burrow of a potential prey 
item (crab, skink or bird). The number of foraging 
burrows was totalled each month. Occupied burrows, 
and those with tracks appearing from the entrance 
after smoothing, not identified as foraging burrows, 
were classified as retreat burrows. The number of 
retreat burrows observed was totalled monthly. 

A temperature data logger (model 6003 A, Unidata 
Australia, Perth) was placed on top of one of the 
dunes with two probes extruding from the logger's 
case. One of the probes was placed 1 cm below the 
sand surface, while the other was buried at 50 cm 



■y ( - 



' ~/' 



gpj 



I 



*% 



Figure 2. Burrow of Varanus panoptes . 

below the surface. A digital temperature reading was 
taken by the logger every 30 minutes. The data were 
downloaded and compiled at the end of every month. 

Correlation analysis was done between the num- 
ber of foraging and retreat burrows counted each 
month and the mean burrow depth. Correlation analy- 
sis was also done between the number of foraging 
burrows, retreat burrows and burrow depths and 
monthly maximum and minimum sand temperature at 
the surface and at 50 cm depth. 

Results 

Overall 93 burrows were identified as either retreat or 
foraging burrows. Fifty-one were identified as forag- 
ing burrows and 42 as retreat burrows, three of the 
retreat burrows were identified as such because they 
were occupied by a monitor lizard. The numbers of 
retreat and foraging burrows observed each month are 
shown in Figure 3. There was a peak in the number of 
retreat burrows in July with a steep drop in August 
and September. The number of foraging burrows 
observed peaked in April, although the number stays 
high until July when the number drops. The number 
of retreat burrows exceeded the number of foraging 
burrows between May and September. 

There was a positive correlation between the number 
of retreat burrows and the burrow depths recorded 
each month (r = 0.67; P = 0.017; Table 1). 

Table 1 . Correlation coefficients between the number 
of retreat and foraging burrows and burrow depths. 

* denotes a significant correlation 



Retreat Burrows 0.67 0.017* 

Foraging Burrows 0.097 0.763 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 27 



12 



10 



O 



3 
SQ 
- 

o 

6 
z 



-2 




O RETREAT BURROWS -Q FORAGING BURROWS 



JAN 



SEP 



MAR MAY JUL 

MONTH 

Figure 3. Monthly number of foraging and retreat burrows of V. panoptes at Fog Bay. 



NOV 



However, the number of foraging burrows is not 
correlated to burrow depths (Table 1 ). The number of 
retreat burrows are negatively correlated with the min- 
imum sand-surface temperatures (r = 0.463; P = 
0.024) and the maximum sand temperature at 50 cm (r 
= -0.616; P = 0.033; Table 2). Burrow depths were 
also negatively correlated to the minimum sand sur- 
face temperature (r = -0.789; P = 0.002) and the maxi- 
mum sand temperature at 50 cm (r = -0.64; P = 0.033; 
Table 2). The number of foraging burrows was unaf- 
fected by sand surface temperature and sand tempera- 
ture at 50 cm (Table 2). This equates to more and 
deeper retreat burrows being dug, the cooler the sand 
temperature at the surface and at 50 cm. 

Discussion 

The field experiments herein were the first attempt 
with this modified design of burrow viewing equip- 
ment and an assessment of its effectiveness is war- 
ranted to justify the results. The only problem 
encountered was when the original silicon glue (a 
metal sealant), used to hold the camera to the jar cas- 
ing, was suspected of having a corrosive effect on the 
camera. The camera stopped working within a short 
time and, when inspected, small white spots were 
seen on the solder of the circuit board. The camera 
worked properly when the spots were cleaned off and 



a non-corrosive sealant (glass/window sealant) was 
used. From then on, the system provided clear images, 
with effective viewing of monitor lizard presence in 
burrows and the depth, width and curvature of all bur- 
rows. The narrower, rounder burrows of other animals 
were easily identifiable at the bottom of foraging bur- 
rows. 

The number of foraging burrows was independent 
of depth or temperature influences. However, the 
greater the number of retreat burrows dug, the greater 
their depth. The number and depth of retreat burrows 
was also negatively correlated to minimum sand-sur- 
face temperature each month. This is indicative of an 
increased need for the lizards to burrow to an increas- 
ing depth as the overnight temperatures cooled. 

Cooler sand temperatures at 50 cm also caused an 
increase in the number and depth of retreat burrows. 
Since thermal diffusion in sand is slow, sand tempera- 
ture at 50 cm heats and cools slowly (Packard and 
Packard, 1988). The sand temperature at 50 cm at Fog 
Bay was the highest overnight when the sand surface 
is lowest (Guinea, 1994). Thus, monitors may be bur- 
rowing more in the dry season to exploit the warmer 
overnight internal sand temperatures. Maximum inter- 
nal sand temperatures fall during the dry season and 
the monitors therefore dig further to find even warmer 
temperatures. Foraging burrows were also responsible 
for increasing the number of burrows seen in the dry 



Vol. 9, p. 28 



Asiatic Herpetological Research 



2001 



Table 2. Correlation coefficients between the number of retreat and foraging burrows and burrow depth and sand 
surface (T so ) minimum and maximum temperatures and sand at 50 cm depth (T S50 ) minimum and maximum tem- 
peratures. * denotes a significant correlation. 





Retreat burrows 
r P 


Foraging 
r 


burrows 
P 


Burrow 

r 


depth 
P 


Min. T so 
Max. T so 


-0.643 
0.316 


0.024* 
0.317 


0.18 

0.227 


0.576 
0.478 


-0.789 
0.722 


0.002* 
0.008* 


Min. T S50 


-0.296 


0.351 


-0.145 


0.653 


-0.086 


0.79 


Max. T S50 


-0.616 


0.033* 


0.026 


0.937 


-0.646 


0.033* 



season, but their depths were constant, and indepen- 
dent of temperature. 

Of other possible reasons for an increase in moni- 
tor lizard burrowing in the dry season, egg ovipositing 
is unlikely, as no eggs were ever observed within bur- 
rows and V. panoptes. Burrow use to conserve water 
loss has been proven important for some arid zone 
monitor lizards (Green, 1972, Vernet et al., 1988). 
This has not been demonstrated for tropical monitor 
lizards, although there is a considerable difference in 
humidity and rainfall between the wet and dry seasons 
in Australia's wet-dry tropical region (Bureau of 
Meteorology, 1989). Water loss in lizards increases 
with decreasing ambient humidity (Hillman and Gor- 
man 1977) and without physiological adaptations to 
prevent water loss, burrowing to moist sand may be 
utilised to prevent desiccation by V. panoptes when 
ambient humidity decreases. More needs to be investi- 
gated on the water economies, and uses of other water 
sources, in this monitor to determine if water loss is 
influential in increasing burrowing during the dry sea- 
son at Fog Bay. 

Acknowledgments 

I thank Bill Milne, Faculty of Technology, Northern 
Territory University for constructing the burrowscope. 
Greg Hill and Pam Dyer provided feedback on its 
design. Equipment was purchased from Oatley Elec- 
tronics. Project funding was by Australian Geo- 
graphic, The Queen's Trust for Young Achievers and 
The Centre for Tropical Wetlands Management, 
Northern Territory University. 



Literature Cited 

Auffenberg, W. 1983. The burrows of Varanus benga- 
lensis: characteristics and use. Records of the Zoolog- 
ical Survey of India. 80:375-385. 

Bennett, A. F. and K. A. Nagy. 1977. Energy expendi- 
ture in free ranging lizards. Ecology 58:697-700. 
Blamires, S. J. 1999. Quantifying predation on sea 
turtle nests by varanids at Fog Bay. MSc Thesis. 
Northern Territory University. 

Bureau of Meteorology. 1989. Climate of Australia. 
Morphea Press, Canberra. 

Capel, V. 1993. Security Systems and Intruder 
Alarms. Newnes, Oxford. 267 pp. 
Cowles, R. B., and Bogert, C. M. 1944. A preliminary 
study of the thermal requirements of desert reptiles. 
Bulletin of the American Natural History Museum 
83:261-296. 

Dyer, P. K., and G.J.E. Hill. 1991. A solution to the 
problem of determining the occupancy status of 
wedge-tailed shearwater Puffinus pacificus burrows. 
Emu 91:20-25. 

Dyer, P. K., and K. Aldworth.. 1998. The "burrow- 
scope": modifications to burrow viewing equipment. 
Emu 98:143-146. 

Green, B. 1972. Water loss of the sand monitor lizard 
(Varanus gouldii) in its natural environment. Ecology 
53:452-457. 

Guinea, M. L. 1994. A possible model to explain win- 
ter nesting by the flatback turtle, Natator depressus at 
Fog Bay, Northern Territory. Pp. 154- 155. In R. James 
(ed.), Australian Marine Turtle Conservation Work- 
shop. Queensland Department of Environment and 



2001 Asiatic Herpetological Research Vol. 9, p. 29 



Heritage, Australian Nature Conservation Agency. 
Gold Coast. 

Hansell, M. H. 1993. The ecological impact of animal 
nests and burrows. Functional Ecology 7:5-12. 

Hillman, S. S., and G. C. Gorman. 1977. Water loss, 
desiccation tolerance and survival under desiccating 
conditions in two species of Carribean Anolis. Oeco- 
logia 29:105-1 16. 

Packard, G. C, and M. J. Packard. 1988. The physio- 
logical ecology of reptilian eggs and embryos. Pp. 
523-605. In C. Gans and R. B. Huey (eds.). Biology 
of the Replilia, Vol. 16. Alan Liss, New York. 

Pianka, E. R. 1969. Habitat specificity, speciation and 
species density in Australian desert lizards. Ecology 
50:498-502. 

Vernet, R.. M. Lemire. and C. Grenot 1988. Field 
studies on activity and water balance of a desert mon- 
itor Varanus griseus (Reptilia: Varanidae). Journal of 
Arid Environments. 15:81-90. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 30-33 



Notes on the Diet, Survival Rate, and Burrow Specifics of Uromastyx aegyptius 
microlepis from the United Arab Emirates 

Peter L. Cunningham 

P.O. Box 17258, AlAin. United Arab Emirates, email: plc@ emirates. net.ae 

Abstract.- Uromastyx aegyptius microlepis are almost exclusively herbivorous and feed on a variety of plant 
species with Pennisetum divisum being extensively utilized in the United Arab Emirates. The survival rate, as 
determined over a one year period without any rainfall is <50%. Burrow openings are mainly aligned in a 
southerly and easterly direction with the average burrow entrance size being 60x155 cm. Temperatures measured 
at 30 cm down the burrow entrance are on average 6°C lower than the ambient temperature. 



Key words.- Reptilia. Squamata. Agamidae, Uromastyx aegyptius microlepis. United Arab Emirates, ecology 




Figure I.The Spiny-tailed lizard, Uromastyx aegyptius 
microlepis. 

Introduction 

Uromastyx aegyptius microlepis (Spiny-tail Lizards; 
Fig. 1) belong to the Family Agamidae and occur 
throughout the Arabian Peninsula, Iran, Iraq and Jor- 
dan (Arnold 1986. Leviton et al. 1992). They are 
diurnal ground dwelling lizards that may reach sizes 
of up to 60 cm and are generally yellowish-grey in 
colour with an impressive spiny club-like tail. Adult 
U. a. microlepis are documented as being mainly her- 
bivorous, although insects form part of their diet, 
while juvenile lizards are thought to be mainly insec- 
tivorous (Arnold 1984, Baha El Din 1996, Brown 



Figure 2. The study site and habitat of U. a. microlepis 
in Abu Dhabi. 

982, Highveld and Slimani 1998, Jongbloed 1997, 
Manthey and Shuster 1996). 

Material and Methods 

Twenty mature Uromastyx aegyptius microlepis indi- 
viduals in a scattered population, ±35 km northwest of 
Al Ain (24°25'07"N and 55°35'01"E; Fig. 2) in the 
Abu Dhabi Emirate of the United Arab Emirates 
(UAE), were observed for one full day each (sunrise 
to sunset). These observations took place during sum- 
mer (May and June 1999) and winter (December 1999 
and January 2000), respectively. I spent 480 observa- 
tion hours studying these lizards. Direct observations 
of plant species utilized during their feeding bouts 
was conducted using an 8x40 binocular over a dis- 
tance of 100m so as not to influence the feeding 
behavior. Tracks were also followed once the lizards 
had retreated underground to confirm sightings. Fecal 
pellets were collected for later analysis. Burrow 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 31 



information (orientation, burrow entrance height and 
width, temperature above ground and 30 cm down 
burrow) was collected from 25 active burrows once 
the lizards had retreated. 

Survival rate was determined for the 20 study individ- 
uals by investigating their burrows for any signs of 
activity in June 2000, one year after the first observa- 
tions were made. 

Results 

Diet. Uwmastyx aegyptius microlepis utilized the fol- 
lowing 10 plant species: 

Class Monocotyledonae 

Gramineae (Grass family): Pennisetum divisum, 
Stipagrostis plumosa 

Class Dicotyledonae 



Amaranthaceae (Cockscomb family): Aerva jav- 
anica 

Asclepiadaceae (Milkweed family): Leptadenia 
pyrotechnica 

Boraginaceae (Borage family): Moltkiopsis cili- 
ata, Heliotropium kotschyi 

Chenopodiaceae (Goosefoot family): Haloxylon 
salicomicum 

Cucurbitaceae (Gourd family): Citrullus colocyn- 
this 

Leguminosae (Pea family): Tavemiera cuneifolia 

Polygalaceae (Milkwort family): Polygala eri- 
optera 

Survival Rate. Of the 20 study individuals first 
observed in May 1999, only 9 were still present in 
June 2000. 



Table 1 . Vegetation selected by Uromastyx aegyptius microlepis as documented by different authors from the 
Arabian Peninsula. * Pulicaria glutinosa observed being utilized in other areas although not in the present study 
area. 



This Study 
UAE- Abu Dhabi 

Aerva javanica 



Citrullus colocynthis 

Haloxylon salicomicum 
Heliotropium kotschyi 



Leptadenia pyrotechnica 
Moltkiopsis ciliata 

Pennisetum divisum capitata 

Polygala erioptera 
*Pulicaria glutinosa 
Stipagrostis plumosa 
Tavemiera cuneifolia 



Jongbloed(1997) 
UAE - Sharjah 



Fagonia sp. 



Pennisetum divisum 



Mandevile(1965) 
Saudi Arabia 



Aristida plumosa 
Astragalus gyzensis 
Citrullus colocxnthis 



Horwoodia dicks on eae 
Launaea capitata 

Moltkiopsis ciliata 
Neurada procumbens 

Plantago boissieri 



Zygophyllum sp. 



Vol. 9, p. 32 



Asiatic Herpetological Research 



2001 



Burrow specifics (n=25). Burrow openings were ori- 
entated as follows: 

North 2, West 3, South 12 (S/West: 4, South: 5, S/ 
East: 3) and East 8. 

Mean average burrow opening height and width 
was 61 cm and 155 cm (n=25), respectively. 

Mean average temperature at a depth of 30 cm 
down the burrow was 38°C (n=25). The ambient tem- 
perature measured at 1.5 m above ground at 13h00 
was 44°C. 

Discussion 

Diet. Ten perennial plant species are selected by Uro- 
mastyx aegyptius microlepis as observed during the 
study period, with the coarse desert grasses Pennise- 
tum division and Stipagrostis plumosa being favored. 
Jongbloed (1997) identified 3 plant species selected 
by the lizards from the Sharjah area in the UAE while 
Mandaville (1965) identified 8 plant species from the 
stomach contents of six Uromastyx aegyptius speci- 
mens from Saudi Arabia (See Table 1). Foley et al. 
(1992) identified 23 plant species, mainly annuals, 
selected by the same species in Israel. 

As no rain had fallen during the study period no 
annuals were observed and therefor the exclusion 
from the diet. It would therefor be expected that more 
plant species, especially flowering annuals, would be 
utilized after rains. Rainfall is highly variable 
(± 100mm p. a.) and unpredictable in the UAE (Bot- 
tomley 1996) forcing the lizards to rely on the avail- 
able perennials. According to Zari (1996 and 1998) 
Uromastyx philbyi feed on a variety of annual grasses 
and perennial shrubs in western Saudi Arabia. High- 
field and Slimani (1998) indicate Uromastyx acan- 
thinurus's preference for the family Chenopodia (fat 
hen/spinach) as well as the salt tolerant Artiplex genus 
and a wide variety of ephemeral plants after rains in 
Morocco. 

An analysis of 170 fresh adult fecal pellets (85 
summer and winter, respectively) confirmed the her- 
bivorous nature of U. a.microlepis as only one pellet 
(<1%) contained insect remains, that of a Tennebri- 
onid beetle. According to Highfield and Slimani 
(1998) adult fecal pellets of U. acanthinurus in 
Morocco contained 6% insect matter. Brown (1982) 
also mentions U. a.microlepis feeding on locusts from 
the UAE. It still has to be investigated if the lizards 



would show a preference for insects during autumn 
and spring although it is expected that the lack of 
insects in the diet could be contributed to the fact that 
most insects, Tenebrionidae at least, are crepuscular 
or nocturnal (Tiger 1996). 

Other items consumed by Uromastyx as observed 
during the fecal analysis of the pellets include: sheep 
droppings (17 pellets), date kernels (4 pellets) and 
feathers (3 pellets). It is expected that above men- 
tioned items are consumed erroneously by the lizards 
as insect material although Brown (1982) suggests 
that sheep droppings be consumed to extract undi- 
gested plant material. If the sheep droppings, date 
kernels and feathers were consumed as insect mate- 
rial, it would suggest that the lizards would prey on 
insects more often if they were more readily available. 

Survival rate. Nine (45%) of the 20 study individuals 
survived a period of one year from May 1999 to June 
2000. Three of the 1 1 missing individuals were found 
as severely malnourished carcasses. The other 8 indi- 
viduals not accounted for are presumed to have suc- 
cumbed below ground in their burrows, some of 
which were covered by sand. No signs of emigration 
were determined. No rainfall occurred during this 
period in the study area. The area is also frequented 
by large numbers of domestic camels and sheep, 
which compete directly with the lizards for the avail- 
able food. It is suggested that the lack of rainfall and 
lack of annual plants as a result of this, together with 
the extra pressure of domesticated stock, resulted in 
the high mortality observed. 

Burrow specfics. Burrow openings were mainly 
aligned to the south and east with 80% of the burrow 
entranced facing this way. During summer, tempera- 
tures consistently reach between 40°C and 47°C 
between lOhOO and 16h00. North and west facing 
burrow entrances would be hotter than burrows facing 
in other directions, especially during midday. During 
winter the lizards would also benefit from the south- 
erly orientation warming the burrows quicker and 
more effectively. Wind direction during summer is 
predominantly from the northwest and can cause 
severe sandstorms locally known as "shamal" (El- 
Ghonemy 1985). This causes lizard whose burrows 
face that direction to spend more time on den clearing, 
a time consuming an energetically costly affair espe- 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 33 



cially during summer when plant availability and pal- 
atability are low. 

Burrow size is often related to the size of the resi- 
dent lizard. The mean average size of 61 x 155 cm 
(n=25) is indicative of an adult Uromastyx colony. 
Bigger burrow openings have higher temperatures at a 
depth of 30 cm, especially when facing to the north or 
west. 

The mean average temperature of 38°C, during the 
heat of the day, at a depth of 30 cm down the burrow 
is on average 6°C lower than the ambient temperature. 
This gradient is imperative to the survival of the liz- 
ards. During the fieldwork it was often found that the 
lizards were resting in the burrows at a depth of just 
over 30 cm, scurrying deeper once the thermometer 
was inserted down the burrow. According to Brown 
(1982), the burrows can be more than 6 feet long and 
3 to 4 feet deep in the form of a spiral. Jongbloed 
(1997) confirms this depth even indicating reports of 
up to 12 feet deep burrows. The same author men- 
tions that an excavated burrow of 5 feet in depth was 2 
feet across and 1 foot high and still not at its end. 
Burrows do not only serve as thermoregulatory 
havens for Uromastyx aegyptius microlepis, but also 
for the Desert Monitor (Varanus griseus) (Pers.obs.) 
and certain Larks (Cunningham 2000, Williams et al. 
1999). Williams et al. (1999), states that Uromastyx 
burrows can potentially reduce Hoopoe Lark (Alae- 
mon alaudipes) water loss by as much as 81% during 
the hottest periods during summer. Temperatures at a 
depth of 30 cm are relatively stable therefor being 
exploited by many desert dwelling animals (Love- 
grove and Knight-Eloff 1988). Burrows are thus an 
effective way of escaping predators, the harsh desert 
environment as well as controlling water loss. 

Literature Cited 

Arnold, E. N. 1984. Ecology of lowland lizards in the 
eastern United Arab Emirates. Journal of Zoology, 
London 204:329-354. 

Arnold, E. N. 1986. A key and annotated checklist to 
the lizards and amphisbaenians of Arabia. Fauna of 
Saudi Arabia 8:385-435. 

Baha El Din, S. 1996. Terrestrial reptiles of Abu 
Dhabi. Pp. 124-147. In P.E. Osborne (ed.), Desert 
Ecology of Abu Dhabi. Pisces Publications, Newbury, 
UK. 

Bottomley, N. 1996. Recent climate of Abu Dhabi. 
Pp. 36-49. In: P.E. Osborne (Editor), Desert Ecology 
of Abu Dhabi. Pisces Publications, Newbury, UK. 



Brown, J. N. B. 1982. Spiny-tailed Agamid - Uro- 
mastyx microlepis (Arabic "Dhub"). Emirates Natural 
History Group Bulletin Nol6:20. 

Cunningham. P. L. 2000. The use of burrows by Hoo- 
poe Lark (Alaemon alaudipes). Tribulus 10. 1 :2 1 . 

El-Ghonemy, A. A. 1985. Ecology and flora of Al Ain 
Region. University of the United Arab Emirates, Al 
Ain. 

Foley. W. J., A. Bouskila, A. Schkolnik and I. Chosh- 
niak. 1992. Microbial digestion in the herbivorous liz- 
ard Uromastyx aegyptius (Agamidae). Journal of 
Zoology, London. 226:387-398. 

Highveld, A. C. and T. Slimani. 1998. The Spiny- 
Tailed Lizard at home: Uromastyx acanthinurus in 
Southern Morocco. Reptiles - Guide to keeping rep- 
tiles and amphibians, July 1998: 76-87. 

Jongbloed, M. 1997. Observations in a Dhub colony. 
Tribulus 7.2:23-24. 

Leviton, A. E., S. C. Anderson, K. Adler and S.A. 
Minton. 1992. Handbook to Middle East Amphibians 
and Reptiles. Scociety for the Study of Amphibians 
and Reptiles, Oxford, Ohio. 252 pp. 

Lovegrove, B. G. and A. Knight-Eloff. 1988. Soil and 
burrow temperatures, and the resource characteristics 
of the social mole-rat Cryptomys damarensis (Bathy- 
ergidae) in the Kalahari Desert. Journal of Zoology, 
London 216:403-413. 

Mandeville, J. 1965. Plants eaten by Uromastyx 
microlepis Blanford and other notes on this lizard in 
Eastern Arabia. Journal Bombay Natural History 
Society 62(1): 161 -163. 

Manthey, U. and N. Schuster. 1996. Agamid Lizards. 
T.F.H. Publications Inc., USA. 

Tiger, B. J. 1996. A preliminary assessment of the 
arthropods of Abu Dhabi. Pp. 172-195. In P.E. 
Osborne (ed.). Desert Ecology of Abu Dhabi. Pisces 
Publications, Newbury, UK. 

Williams, J. B., B. I. Tielman and M. Shobrak. 1999. 
Lizard burrows provide thermal refugia for larks in 
the Arabian Desert. The Condor 101 (3):7 14-7 1 7. 

Zari, T. A. 1996. Effects of body mass and tempera- 
ture on standard metabolic rate of the herbivorous 
desert lizard, Uromastyx philbyi. Journal of Arid Envi- 
ronment 33:457-461. 

Zari, T. A. 1998. Effects of sexual condition on food 
consumption and temperature selection in the herbiv- 
orous desert lizard, Uromastyx philbyi. Journal of 
Arid Environment 38:371-377. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 34-70 



The Amphibians and Reptiles of Panay Island, Philippines 

John W. Ferner 1, 2 , Rafe M. Brown 2, 3 , Rogelio V. Sison 4 and Robert S. Kennedy 2 

Department of Biology, Thomas More College, Crestview Hills, Kentucky 41017, USA. email: 

fernerj@thomasmore.edu; Geier Collections and Research Center, Museum of Natural History and Science, 

1301 Western Avenue, Cincinnati, Ohio 45203, USA; Section of Integrative Biology and Texas Memorial 

Museum, University of Texas, Austin, Texas 78712, USA. email: rafe@mail.utexas.edu; 4 Zoology Division, 

National Museum of the Philippines, Executive House, P. Burgos Street, Manila, Philippines, email: 

nmzoo@fastmail.i-next.net. 

Abstract.- We provide species accounts for 20 amphibians (frogs and toads) and 72 reptiles (one turtle, 36 lizards, 
and 35 snakes) from the central Philippine island of Panay and its associated land-bridge islets. Panay is a 
moderately-sized island (123,000 km - ) that currently is separated from the nearby islands of Negros, Masbate, 
and Cebu by shallow seas, indicating that dry land connections once allowed free exchange of flora and fauna 
between these land masses at several periods during the Pleistocene. This fact, coupled with the wealth of 
knowledge on herpetological communities of Negros and Cebu, has led biologists to assume that the amphibians 
and reptiles of Panay are reasonably well known. Our data suggest that this is far from true. Our recent survey 
work (1989 until present) has resulted in the discovery of at least six (and probably 12-15) new species of 
vertebrates, most of which appear to be high elevation rainforest Panay endemics. In this paper we note 
numerous taxonomic problems that await the attention of biologists; these surely will result in an increase of the 
known species from Panay. We expect that many widespread species complexes currently of uncertain taxonomic 
status will eventually be recognized as Panay endemics, further bolstering the total diversity and degree of 
endemism on Panay. The status of Panay herpetofauna warrants particular attention because many of the 
suspected new and endemic species appear to be forest obligates. With the near complete removal of Panay 's low 
elevation forests and increased pressure on its mid- and upper montane forests, we suspect that many of Panay's 
endemics are threatened by extinction. Panay should be regarded as a tropical island with a unique geological 
history, a spectacular compliment of habitat types, and a diverse set of species communities - all of which are 
beset with a conservation crisis of deforestation and probable extinction. Panay may be a particularly appropriate 
model island for large scale conservation efforts aimed at sustainable management of forest resources. However, 
before informed conservation and management plans can be enacted, additional biodiversity survey work must 
be conducted on Panay. 

Key words.- Reptiles, amphibians, SE Asia, Philippines, Panay, Visayan Islands, conservation crisis, 
herpetofauna. 



Introduction 

The central (or "Visayan") Philippine island of Panay 
(Figs. 1 , 2), is part of a Pleistocene aggregate island 
complex that includes Negros, Cebu, Guimaras, Mas- 
bate, Ticao, and several small land-bridge islands 
associated with each of these larger land masses. 
Although amphibian and reptile species similarity 
among Visayan Islands was noted much earlier (Tay- 
lor, 1920, 1922a-d, 1928; Inger, 1954; Leviton, 
1963c; Brown and Alcala, 1970a), it was not until 
Heaney (1985, 1986) explicitly illustrated the position 
of the 120 m underwater bathy metric contour in the 
Philippines that the land bridge implications of chan- 
nel depth became fully appreciated by biologists. It is 



now understood that five major Philippine island 
groups, (complexes of islands separated by shallow 
channels) intermittently formed much larger land 
mass amalgamations at various times during the mid- 
to late-Pleistocene (Fig. 1). It is presumed that these 
events allowed free exchange of fauna and flora via 
land-positive connections between the otherwise iso- 
lated islands of today. Each of the Philippine Pleis- 
tocene aggregate island platforms (Fig. 1) are now 
recognized by biogeographers as subprovinces (Tay- 
lor, 1928) due to the fact that each supports highly- 
celebrated suites of endemic taxa (Taylor, 1928; Dick- 
inson, 1991; Heaney and Regalado, 1998; Heaney et 
al., 1998; Alcala and Brown, 1998; Kennedy et al., 
2000). 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 35 




Figure I.The Philippines (darkly shaded islands), with 
the major Pleistocene aggregate island platforms indi- 
cated by tracing of the 120 m submarine bathymetric 
contour (following Heaney, 1985, 1986). Star = Manilla. 



One unfortunate result of the prevailing perspec- 
tive since Heaney 's (1985) landmark paper, has been 
herpetologists' lack of attention to islands that are 
land-bridge (once connected by dry land) to well sur- 
veyed members of the same Pleistocene island com- 
plex. A case in point is Panay, a central Philippine 

island of 123,000 km" with several mountain peaks of 
more than 2000 m in elevation (Figs. 1, 2). Perhaps 
not unreasonably, numerous biologists over the past 
century have assumed or suggested that a species' 
presence on the well-surveyed Negros implied its 
undocumented presence on Panay as well (see Inger, 
1954; Leviton, 1963; Brown and Alcala, 1970, 1978, 
1980, 1986; Brown and Rabor, 1967; Alcala, 1986). 
Nevertheless, cases of low-level island endemism 
have been documented on individual islands within 
Pleistocene island complexes (Alcala, 1958, 1962; 
Taylor, 1920; 1922a-d; 1923; 1925; Inger, 1954; 



0-150 

150-400 

400-800 

800-1200 

1200-1600 

> 1600 



Figure 2. Panay Island, its position in the Philippines indicated by darkened arrow (inset). Discrete shading indi- 
cates increasing elevational increments (see key), enclosed numbers indicate collection localities (see Table 1 ), 
underlined bold text indicates provinces (boundaries indicated with darkened dashed lines), and darkened circles 
indicate major cities. 



Vol. 9, p. 36 



Asiatic Herpetological Research 



2001 



Table 1 . Study sites and collection localities for museum specimens of amphibians and reptiles from Panay Island. 
"No further data available" indicates specimens for which municipalities were the only available collection data in 
museum catalogs. The term "municipality" is synonymous with "town" but municipalities are also the political 
entity surrounding towns by these names. Thus, municipalities contain small surrounding villages (Barrios 
[="Barangays "] which may contain smaller "Sitios"). It is reasonable to assume that most collections localized to 
the municipality level were also collected in the town of the same name. Entries including "and vicinity" indicates 
specimens that may have been collected in the municipal town, or they may be from smaller surrounding Baran- 
gays. * Combined into a single locality due to the close proximity of collection sites at the common borders of 
these municipalities. 



Site 



Province 



Municipality 


Notes 


Ibajay 


Barangay Bugtong-bato 


Libacao 


Libacao town "and vicinity" 


Makato 


Makato town "and vicinity" 


Malinao 


no specific locality data 


Malay 


Nogas Point 


Nabas 


Nabas town "and vicinity" 


Tangalan 


Tangalan town "and vicinity" 


Malay 


Borocay Isl., no specific locality data 


Barbaza 


Barbaza town "and vicinity" 


Bugasong 


Bugasong town "and vicinity" 


Culasi 


Barangay Alojipan (Mt. Madja-as; 
PNM/CMNH site) 


Tobias Fornier 


Barangay Tobias, Villaflor 


Valderrama 


Barangay San Agustin, (Mt. Baloy and 
vicinity; PNM/CMNH site) 


San Remegio 


Barangay Aningalan, Aningalan 
mountain range 


Tibiao 


no specific locality data 


Caluya 


Caluya Isl., no specific locality data 


Sibay 


Sibay Isl., no specific locality data 


Caluya 


Semirara Isl., no specific locality data 


Hamtik 


Barangay Gunisang-an 


Sibalom 


Barangay Egaiia 


San Jose 


Barangay Bagumbayan 


Capiz 


no specific locality data 


Marubusao 


Burias 


lloilo 


no specific locality data 


lloilo City 


lloilo City "and vicinity" 


Ajuy 


no specific locality data 


Calinog 


Mt. Baloy and vicinity 



1. 


Aklan 


2. 


Aklan 


3. 


Aklan 


4. 


Aklan 


5. 


Aklan 


6. 


Aklan 


7. 


Aklan 


8. 


Aklan 


9. 


Antique 


10. 


Antique 


11. 


Antique 


12. 


Antique 


13. 


Antique 



14. 



Antique 



15. 


Antique 


16. 


Antique 


17. 


Antique 


18. 


Antique 


19.* 


Antique 



20. 


Capiz 


21. 


Capiz 


22. 


lloilo 


23. 


lloilo 


24. 


lloilo 


25. 


lloilo 



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Asiatic Herpetological Research 



Vol. 9, p. 37 





Site 


Province 




Municipality 


Notes 


26. 


lloilo 






Barotac Viejo 


Barangay San Francisco 


27. 


lloilo 






Carles 


no specific locality data 


28. 


lloilo 






Carles 


Sicogon Island; including Buaya area 


29. 


lloilo 






Dingle 


Bulabog-Putian National Park; PNM/ 
CMNH Site 


30. 


lloilo 






Estancia 


no specific locality data 


31. 


lloilo 






Lambunao 


no specific locality data 


32. 


lloilo 






Mandurriao 


no specific locality data 


33. 


lloilo 






Pototan 


Pototan town "and vicinity" 


34. 


lloilo 






Cabatuan 


no specific locality data 


35. 


lloilo 






Estancia 


Gigante North Isl., no specific locality 
data 


36. 


lloilo 






Estancia 


Gigante South Isl., Gabi area 


37. 


lloilo 






Estancia 


Gigante South Isl., no specific locality 
data 


38. 


lloilo 






Calinog 


E. Catalbac ("Calinog town") 


39. 


lloilo 






Estancia 


Calagna-an Isl.; Barangkalan and 
vicinity 


40. 


lloilo 






Concepcion 


ca 1 km NW SBS lyang Beach Resort 


41. 


lloilo 






Concepcion 


Pan de Azucar Isl., no specific locality 
data 


42. 


Negros Occidental 


Pulupandan 


Inampulagan Isl., 8 km W Munic. W. 
Pulupandan, Negros Occ. Prov. 


43. 


lloilo, 


Subprov. 


Guimaras 


Unknown 


Guimaras Isl., no specific locality data 


44. 


lloilo, 


Subprov. 


Guimaras 


Jordan 


Guimaras Isl.; Jordan town "and 
vicinity" 


45. 


lloilo, 


Subprov. 


Guimaras 


Unknown 


Guimaras Isl.; 2.1 km SW Daan 
Banwa 


46. 


lloilo, 


Subprov. 


Guimaras 


Nueva Valencia 


Guimaras Isl.; Nueva Valencia town 
"and vicinity" 


47. 


lloilo, 


Subprov. 


Guimaras 


Buenavista 


Guimaras Isl.; Buenavista town 


48. 


lloilo, 


Subprov. 


Guimaras 


Nueva Valencia 


Panobolon Isl.; Nueva Valencia town. 


49. 


Unknown 




Unknown 


"Masaya" (not on available maps) 


50. 


Unknown 




Unknown 


Panay Island: no Further Data 



Vol. 9, p. 38 



Asiatic Herpetological Research 



2001 



Brown and Alcala, 1961, 1970a-b, 1978, 1980, 
1982a, 1986, 1994; Brown et al, 1997a-b; Brown et 
al., 1999), suggesting that the two large islands war- 
rant individual attention by biogeographers. Despite 
the assumption that many widespread Philippine spe- 
cies were present on Panay (e.g., see Alcala, 1986), 
biogeographical summaries and taxonomic reviews 
historically have listed only nine vouchered (based on 
museum specimens) species of snakes, six geckos, six 
skinks, and seven frogs (Inger, 1954; Leviton, 1963c; 
Brown and Alcala, 1970, 1978, 1980) - 30% of the 
herpetofauna considered in the present report. 

Until very recently the higher elevation forested 
regions of Panay have not been explored by biolo- 
gists. At present, the results of only a few preliminary 
surveys in a few high elevations localities are avail- 
able (Gonzales and Kennedy, 1990, 1996; Sison et al., 
1995; Ferner et al., 1997; Brown et al., 1997a; 1999). 

The purpose of this paper is to synthesize museum 
records that document the diversity, endemism, status, 
and distribution of the amphibians and reptiles from 
Panay and to report on several recent collections 
resulting from the National Museum of the Philip- 
pines/Cincinnati Museum of Natural History Philip- 
pine Biodiversity Inventory (1989-1994). Another 
primary goal is to draw attention to recently-discov- 
ered endemic species of amphibians and reptiles from 
Panay and to stress the need for immediate faunal sur- 
veys on this tropical island beset by a conservation 
crisis of deforestation. 

Material and Methods 

Study sites and collection locations are presented in 
Table 1 and shown in Figure 2. Additional site and 
habitat information for many TNHC, PNM, CAS, 
FMNH, and CMNH specimens may be found in the 
museum catalogs. The PNM/CMNH Philippine 
Biodiversity Inventory team conducted field studies at 
sites 11 and 13 (Figs. 3-9) and various members of 
the team (particularly RVS and RMB), visited other 
locations (Figs. 10-14) to do more limited surveying. 
We established elevational transects in a variety of 
habitat types (Ruedas et al., 1994, as modified by 
Brown et al. 1995a, 1996, 2000) and utilized standard 
collection and specimen preservation techniques 
(Simmons, 1987; Heyer et al., 1994). Detailed exami- 
nation of all material was conducted by RMB and 
JWF. 

We follow the taxonomy of Taylor (1922c), Brown 
and Alcala (1978, 1980) and Brown et al. (1995a; 
1995b) for gekkonid and scincid lizards. The taxon- 
omy of Inger (1954, 1996), Frost (1985), Duellman 



(1993), Brown and Alcala (1994), Inger and Tan 
(1996), Alcala and Brown (1998) and Emerson et al. 
(2000), was consulted for amphibians and we include 
(in parentheses) the unsubstantiated taxonomic 
hypotheses of Dubois (1992) for reference. While no 
suitable taxonomy currently is available for Philippine 
agamids (see Taylor, 1922c), we consulted Musters 
(1983) and McGuire and Alcala (2000) for identifica- 
tion of our Draco specimens. Snake taxonomy was 
based on Taylor (1922a), Leviton's "Contribution to a 
Review of Philippine Snakes" series (Leviton 1961, 
1962, 1963a-c, 1964a-d, 1965, 1967, 1968, 1970a-b, 
1979, 1983, 1985), Leviton and Brown (1959), Inger 
and Marx (1965), Malnate and Underwood (1988), 
Wynn and Leviton (1993), McDiarmid et al. (1999), 
and Brown et al. (1999). Scale counts on snakes were 
taken using the methods of Dowling (1951a, 1951b). 
Field techniques in our surveys have been described 
in Ruedas et al. (1994) and Brown et al. (1995a; 1996; 
2000). Museum abbreviations follow Duellman et al. 
(1978) and Leviton et al. (1985). 

In order to examine large scale faunal similarities 
between Panay and other large island of the Philip- 
pines, we assessed overall herpetofaunal simmilarities 
by calculating coefficients of similarity using a simple 
index (C - 2W/a+b; see Brown and Lomolino, 1998, 
for review) for the amphibians and reptiles of Panay 
and other islands. Our data for these calculations were 
based on all available literature (above) and updated 
conservative estimates of the taxonomy of Philippine 
amphibians and reptiles (Brown, Crombie, Diesmos, 
unpublished data). 

Results 

We present records for 20 amphibians (anurans; Figs. 
15-25), and 72 reptiles (one turtle, 36 lizards, and 35 
snakes; Figs 26-52) from Panay and its associated 
land-bridge islets. The results of faunal similarity cal- 
culations are presented in Figure 53. Individual spe- 
cies accounts, with comments on their status, are 
presented below. 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 39 



Species Accounts 

Amphibia 

Anura 

Bufonidae 

Bufo marinus (Linnaeus) (Fig. 15) 

Introduced on most inhabited islands of the Philip- 
pines (Inger, 1954; Diesmos, 1998; Alcala and Brown, 
1998), this species is particularly common in agricul- 
tural areas and near human habitation. It is wide- 
spread and abundant in disturbed and agricultural 
areas on Panay. 

Localities and specimens: (Site 1) PNM 1 144 — 47, 

1228-33; (Site 11) CMNH 4958-59, PNM 1289-96, 
2552-54; (Site 12) PNM 1232-33; (Site 19) CMNH 
5197; (Site 23) USNM 339985-86; (Site 24) FMNH 
61482-84; (Site 31) PNM 1 144-47; (Site 33) USNM 
339987-88, 340062. 

Microhylidae 

Kaloula conjuncta negrosensis Taylor (Fig. 
16) 

Taylor (1920, 1922a) first collected this form on 
Negros and Guimaras islands. He (Taylor, 1922a) rec- 
ognized it as a full species but Inger (1954) placed K. 
negrosensis in the synonymy of Luzon's Kaloula con- 
juncta; no further taxonomic studies have been forth- 
coming. Kaloula conjuncta negrosensis is both a 
burrower and a climber (Taylor, 1920; Alcala, 1962, 
1986); it has been found in the detritus of the forest 
floor (750-1075 m on Negros; Alcala, 1962) and in 
the leaf axils and holes of trees (Alcala 1962). They 
also emerge and congregate around water that collects 
in pools and ditches in the rainy season (Taylor, 1920, 
1922a). Our Mt. Madja-as specimen was found in a 
tree hole 10 cm from the ground (tree < 0.5 m dbh). 
Specimens from near the base of Mt. Baloy were 
found in breeding aggregations around water buffalo 
wallows near a large river. We suspect that the 
Visayan populations represent a distinct evolutionary 
lineage and that they will eventually once again be 
recognized as a full species once new data become 
available (Brown and Diesmos, unpublished data). 

Localities and specimens: (Site 8) CAS 127890; 
(Site 1 1) PNM 2555; (Site 13)TNHC 56340-46; (Site 
16) CAS 127510-11, 127591; (Site 18) CAS 127815; 
(Site 47) CAS 124446. 



Kaloula picta (Dumeril and Bibron) (Fig. 17) 

This frog is found at low elevations (100-200 m) in 
open areas near human habitations (Boulenger, 1882; 
Alcala, 1956, 1958). It is a burrowing species that 
conceals itself under leaf litter and topsoil until the 
beginning of the rainy season. Choruses may contain 
hundreds of individuals (Alcala, 1962; Alcala and 
Brown, 1998). This appears to be the first published 
account of this species on Panay. 

Localities and specimens: . (Site 16) CAS 127617- 
37; (Site 18) CAS 127816, 127827; (Site 23) CAS- 
SU 14219-20; USNM 78079-80, 78842. 

Kaloula sp. 

In 1992, while conducting survey work at Barangay 
Alojipan (Site 11), we heard the distinctive honking 
calls of a forest species of Kaloula. Although we were 
unable to locate and collect specimens, we noted that 
the calls were superficially similar to Kaloula kalin- 
gensis (Taylor, 1922a; single honk, delivered approxi- 
mately every 15 to 20 minutes) from Luzon. Neither 
Kaloula kalingensis nor the related K. kokacii (Ross 
and Gonzales, 1991) have been reported from the 
nearby island of Visayas, although recent field work in 
the last remaining low elevation forests of Negros 
(Municipality of Ayungon, Negros Oriental Prov.; C. 
N. Dolino and A. C. Diesmos, personal communica- 
tion) reveals the presence of a forest species (also pre- 
viously unrecorded) with single honking calls there as 
well. Gaulke (in press) recently has discovered a pop- 
ulation of Kaloula in NW Panay that may be the same 
as that heard (but not collected) previously on Panay 
and Negros. 

Localities and specimens: none. 

Ranidae 

Limnonectesc\. leytensis (Boettger) (Fig. 18) 

This species is widely distributed in patches and con- 
sidered common in some localities on the nearby 
Negros (Alcala, 1986; Alcala and Brown, 1998). 
However, it has been collected from only one locality 
on Panay (Sison et al., 1995). On Negros, this species 
inhabits coolor high elevation mountain streams 
between 150 and 900 m (Alcala, 1962) and probably 
breeds terrestrially but deposits eggs in close proxim- 
ity of water (Alcala and Brown, 1956; Alcala, 1962; 
Brown and Alcala, 1982b). When hatching, terrestrial 
embryos of these and related species fall, are carried 
by males, or are washed into water (Alcala, 1962; see 
also Inger et al., 1986; Brown and Iskandar, 2000). 
We find it doubtful that Visayan specimens identified 



Vol. 9, p. 40 



Asiatic Herpetological Research 



2001 



Table 2. List of known species from Panay and smaller, nearby, land-bridge islands and other islands within the 
political boundaries of major Panay Island provinces. Entries include Panay species with vouchered specimens in 
major museum collections (see text and species accounts for discussion of taxonomy used) and the first published 
accounts by authorities utilizing museum specimens. IR = island record or first published record from within major 
Panay island provinces. (1 Previously reported from Panay, nearby islets, or Visayan sea by Alcala (1986) but with- 
out specific reference to specimens. 2 See also Gaulke (in press). 3 Recorded from land-bridge islets but currently 
not recorded from the mainland of Panay. 4 A record from Semirara Isl., within the political boundary of Panay's 
Antique Province, but land-bridge to Mindoro Isl.; not likely to be discovered on Panay in the future. 5 Apparent 
major range extension, in need of verification or based on specimens with locality data probably in error.) 



Bufo marinus 

Kaloula conjuncta negrosensis 
Kaloula picta 
Kaloula sp. 

Limnonectes d leytensis 
Limnonectes visayanus 
Occidozyga laevis 
Platymantis corrugatus 
Platymantis dorsalis 
Platymantis negrosensis 
Platymantis insula tus 
Platymantis panay en sis 
Platymantis sp. 1 
Platymantis sp. 2 
Platymantis sp. 3 
Rana cancrivora cancrivora 
Rana erythraea 
Ranad. everetti 
Rana vittigera 
Po/ypedates leucomystax 
Cuora amboinensis amboinensis 
Bronchocela sp. 
Draco spilopterus 
Hydrosaurus pus tula tus 
Gonocephalus sp. 
Cosymbotus platyurus 
Cyrtodactylus annulatus 



Inger, 1954 

IR (see Taylor, 1920; 1922a) 1 

IR 

IR 2 

IR 

Inger, 1954 

Inger, 1954 

IR 2 

IR 2 

Sisonet al., 1995 

Brown and Alcala, 1970b 3 

Taylor, 1920; Inger, 1954; Brown et al., 1997a 

IR 

IR 

IR 

Inger, 1954 

Inger, 1954 

Sison etal., 1995 

IR 

Inger, 1954 

Gaulke and Fritz, 1998 

Taylor, 1922c 

Musters, 1983 

IR 2 

Sison etal., 1995 

Brown and Alcala, 1978 

IR 2 



2001 



Asiatic Herpetological Research 



Vol.9, p. 41 



Cyrtodactylus philippinicus 
Gehyra mutilata 
Gekko gecko 
Gekko gigante 
Gekko mindorensis 
Hemidactylus frenatus 
Hemidactylus stejnegeri 
Hemiphyllodactylus insularis 
Lepidodactylus lugubris 
L epidodactylus planicaudus 
Brachymeles boulengeri taylori 
Brachymeles talinis 
Brachymeles tridactylus 
Dasia grisea 
Das/a semicincta 
Emoia atrocostata 
Lamprolepis smaragdina philippinica 
Lipinia pulchella taylori 
Mabuya indeprensa 
Mabuya multicarinata borealis 
Mabuya multifasciata 
Parvoscincus sisoni 
Sphenomorphus arborens 
Sphenomorphus coxi divergens 
Sphenomorphus cumingi 
Sphenomorphus jagori grandis 
Sphenomorphus steerei 
Tropidophorus grayi 
Varanus sal va tor nuchal is 
Acrochordus granulatus 
Python reticulatus 
Ahaetulla prasina preocularis 
Boiga angulata 



IR 

Brown and Alcala, 1978 

Taylor, 1922c; Brown and Alcala, 1978 

Brown and Alcala, 1978 3 

Sison et al., 1995 

Brown and Alcala, 1978 

Sison etal., 1995 

Brown and Alcala, 1978 

Brown and Alcala, 1978 3 

Brown and Alcala, 1978 

IR 

Brown and Alcala, 1980 

Brown and Alcala, 1980 

Brown and Alcala, 1980 4 

IR 5 

IR 3 

Brown and Alcala, 1980 

IR 

Sison etal., 1995 

IR 

IR 2 

Ferneretal., 1997 

Brown and Alcala, 1980 

IR 5 

IR 5 

Brown and Alcala, 1980 

Brown and Alcala, 1980 

Sison etal., 1995 

Gaulke, 1991a, 1991b, 1992 

IR 

Leviton, 1963c 

Leviton, 1963c, 1968 

IR 2 



Vol. 9, p. 42 



Asiatic Herpetological Research 



2001 



Bo/gad cynodon 
Bo/ga ci. dendrophila 
Calamaria gervaisi 
Cerberus rynchops 
Chrysopelea paradisi 
Cyclocorus lineatus alcalai 
Dendrelaphis caudolineatus terrificus 
Dendrelaphis pictus p ictus 
Elaphe erythrura psephenoura 
Gonyosoma oxycephala 
Hologerrhum dermali 

Lycodon aulicus capucinus 
Oligodon modestum 
Psammodynastes pulverulentus 
Pseudorabdion mcnamarae 
Pseudorabdion oxycephalum 
Pseudorabdion talonuran 
Tropidonophis negrosensis 
Zaocys luzonensis 
Calliophis calligaster gemianulis 
Hydrophis belcheri 
Hydrophis cyanocinctus 
Hydrophis elegans 
Hydrophis inornatus 
Lapemis hardwickii 
Laticauda colubrina 
Ramphotyphlops braminus 
Rhamphotyphlops cumingii 
Typhlops castanotus 
Typhlops hypogius ( = T. ruber ? ) 
Tropidolaemusc\ wagleri 
Trimereserus flavomaculatus 



IR Z 

IR 

Leviton, 1963c; Ingerand Marx, 1965 

Gyi, 1970 

Sisonetal., 1995 

IR 

Leviton, 1970b 

Leviton, 1963c, 1970b 

Leviton, 1979 

IR 2 

IR 2 

Leviton, 1965 

Sisonietal., 1995 

IR 2 

Sisonietal., 1995 

IR 

Brown etal., 1999 

Leviton, 1963c; Malnate and Underwood, 1988 

Ross etal., 1987 

Leviton 1963b, 1963c 

IR 1 

IR 1 

IR 

IR 

IR 1 

IR 

IR 

IR 

Wynn and Leviton, 1 993 

IR? (see McDiarmid et al., 1999) 

IR 

Gaulke (in press) 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 43 



as Rana cf leytensis are conspecific with specimens 
referred to this species from the Mindanao Aggregate 
Island Complex (Leyte, Samar, Bohol, and Mind- 
anao). 

Locality and specimens: . (Site 25) PNM 1 1 14-15. 

Limnonectes visayanus (Inger) (Fig. 19) 

This large fanged frog is found in clear forest streams; 
it is known to hide in rock crevices during the day and 
call from rocks and stream banks above water at night 
(Alcala, 1962; Alcala and Brown, 1998). We found L. 
visayanus at low elevations on rocks in large rivers 
near Mt. Madja-as. This species may breed and lay 
eggs outside of water (Alcala, 1962). 

Localities and specimens: (Site 2) PNM 1715-20; 
(Site 3) CAS 137592-95, 137590-91, 139164-66, 
USNM 305671-76; (Site 4) PNM 1613-20, 1623-27; 
(Site 6) CAS 137596-98; (Site 7) PNM 1799-800, 
1828-31, 1836-39, 1845, 1855-60, 1865-77; (Site 
1 1 ) CMNH 4894-98, 4899, PNM 1 302-06, 26 17-21; 
(Site 13) TNHC 56337; (Site 14) PNM 3710-12, 
3732, 3764-68, 3805; (Site 25) 1085-92, 1133-38, 
1140; (Site 28) CAS 124093-106, 124442-14, 
124950-58; (Site 39) CAS 124121, 124293-97; (Site 
44) CAS 125308-309, 125312; (Site 47) CAS 
125302-307; (Site 50) USNM 78072-78. 

Occidozyga laevis (Gunther) (Fig. 20) 

Occidozyga laevis is found in flooded fields in agri- 
cultural areas, in road-side ditches and open sewers, 
and in streams and rivers from lowlands to high eleva- 
tion forested sites (Inger, 1954; Alcala, 1962; pers. 
obs.). On Negros this species has an altitudnal range 
of sea level to 1 150 m (Alcala, 1962). Specimens are 
common in stream-side pools along larger rivers in 
forested areas near Mt. Madja-as and Mt. Baloy. 

Localities and specimens: . (Site 1 ) PNM 1 1 10-11, 
1113, 1116-29, 1141, 1184-99; (Site 2) PNM 1116- 
29, 1690-1714, 1721-23, 1731, 1757-58; (Site 3) 
CAS 137586-88, 139148, 139167-68, USNM 
305647-48; (Site 4) PNM 1600-02, 1611, 1621-22; 
(Site 6) CAS 137614-15, USNM 305649, 305650- 
54; (Site 7) PNM 1832-35, 1841-44, 1848-54, 1862- 
64, 1880; (Site 9) PNM 1156, 1165-68, 1170-71; 
(Site 10) PNM 1110-11, 1113; (Site 11) CMNH 
4951-57, PNM 1329, 2655-61; (Site 12) PNM 1 163- 
64; (Site 14) PNM 3730-31, 3782, 3804; (Site 22) 
CAS-SU 9813; (Site 23) CAS-SU 14224-25, 14373; 
(Site 24) FMNH 61478-81; (Site 25) PNM 1141; 
(Site 26) PNM 1160-62; (Site 27) CAS-SU 14049; 
(Site 28) CAS 124959-70, 124059-76, 124426, 
124432-33, 124439; (Site 30) CAS-SU 14223; (Site 



32) PNM 1172-73; (Site 38) CAS 132880, 132887- 
901, 134089-96; (Site 39) CAS 124171, 124190-91; 
(Site 40) USNM 339989; (Site 41) CAS 125001, 
124177, 124184-85, 124194-96; (Site 42) CAS-SU 
23946-49. 23952-58, 23961-63; (Site 44) CAS 
125361-62; (Site 47) CAS 125311, 125344-59. 

Platymantis corrugatus (Dumeril) (Fig. 21) 

This widely distributed terrestrial frog inhabits the 
forest floor from sea level to above 1300 m (Alcala, 
1986). On Mt. Madja-as we found P. corrugatus in 
leaf litter and in limestone crevices. This account and 
that of Gaulke (in press) appear to be the first pub- 
lished records of this species from Panay. 

Localities and specimens: (Site 6) CAS 137616-19, 
139149, 185494; (Site 11) CMNH 4960-63, 5118, 
PNM 2556-59; (Site 13) CMNH 3160-65; (Site 25) 
PNM 1 103-06; (Site 28) CAS 124058. 

Platymantis dorsalis (Dumeril) 

This common forest frog is found in the detritus of the 
forest floor as well as in tree cavities and low tree 
ferns (Alcala, 1962; Alcala and Brown, 1998). 
Although recent studies (Brown et al., 1999) indicate 
the presence of numerous cryptic species in the P. 
dorsalis complex on Luzon (and we suspect that fur- 
ther new species await discovery in the Visayan 
islands), the calls of some P. dorsalis have been heard 
on Mt. Baloy and Mt. Madja-as are, at least superfi- 
cially, similar to the short, whistling, ascending fre- 
quency sweep of true P. dorsalis from Luzon Island 
(Brown et al., 1997c). Thus, while we expect that 
more species in the dorsalis complex will soon be dis- 
covered in the Visayas, we can confidently assert that 
at least some Panay populations are indistinguishable 
from P. dorsalis of Luzon Island (Brown et al., 1997c; 
1999). This account and that of Gaulke (in press) 
appear to be the first published records of this species 
from Panay. 

Localities and specimens: . (Site 2) PNM 1734—56; 
(Site 6) CAS 137620-40, 137649-51, 139150-63, 
USNM 305655-70; (Site 1 1 ) CMNH 4964-98, 5206, 
PNM 2559, 2562-88; (Site 13) TNHC 56347-50; 
(Site 14) PNM 3713-19, 3729, 3733^45, 3756-62, 
3772-74, 3783-85, 3788-89, 3791-96, 3830-31, 
3857, 3860, 3862-65, 3882-83, 3886-88, 3893-95, 
3906-08; (Site 25) PNM 1093-102; (Site 28) CAS 
124419, 124428-31, 124440-41, 124689-91; (Site 
39) CAS 124123-33, 124146-47; (Site 41) CAS 
124041^12, 124122, 124186-89, 125014-19. 



Vol. 9, p. 44 



Asiatic Herpetological Research 



2001 



Platymantis negrosensis Brown, Alcala, 
Alcala, and Diesmos 

This recently-described forest frog (Brown et al, 
1997b) has only been documented from two sites on 
Panay but is also known from localities on the nearby 
island of Negros, from approximately 300 to 1625 m 
in elevation (Alcala, 1958; Alcala and Brown, 1957; 
Alcala, 1962). On that island, this species occupies 
arboreal microhabitats in primary forest (Brown and 
Alcala, 1961; Alcala, 1962; Brown et al, 1997b). We 
documented this island record for Panay (Sison et al., 
1995; then recognized as P. guentheri) from speci- 
mens collected in forest on Mt. Baloy at about 950 m. 
This species is related to P. luzonensis but differs in 
characteristics of the advertisement call an external 
morphology (Alcala and Brown, 1998). 

Localities and specimens: . (Site 13) CMNH 3166; 
(Site 14) PNM 3889. 

Platymantis insulatus Brown and Alcala (Fig. 
22) 

A frog known from primary and secondary forest situ- 
ated on karst limestone outcrops, this species was 
originally discovered (Brown and Alcala 1970b) on 
the forest floor and in the open mouths of small caves 
at low elevations (Alcala and Brown, 1998) on the 
island of Gigante South off Panay's northeast coast. A 
recent (June 2000) visit to the type locality by R. 
Brown and A. Alcala confirmed this species' persis- 
tence despite the complete removal of the original for- 
est. The presence of an endemic species on such a 
small, land-bridge island is puzzling and suggests that 
it may still be (or at least, may have once been) 
present on eastern Panay. Unsurveyed limestone for- 
mations along the northeastern coast are the most 
promising possibility for locating this species on 
Panay. 

Localities and specimens: . (Site 35) CAS 157235- 
39; (Site 36) CAS 137641-12; (Site 37) CAS 
117440-41; 119967-69, MCZA-72946. 

Platymantis panayensis Brown, Brown and 
Alcala 

Only recently described (Brown et al., 1997a) from 
our collections from Mt. Madja-as, this species is 
closely related to P. hazelae from Negros and occu- 
pies similar microhabitats in high elevation cloud for- 
ests. Taylor (1920:101) apparently had a specimen 
(collected by R. McGregor) of this species on hand 
during the description of Philautus (= Platymantis) 
hazelae and he considered it conspecific with the 
Negros population. On the basis of Taylor's (1920) 



account, Inger also (1954) included P. hazelae in his 
list of species from Panay . 

We collected the majority of the type specimens of 
this species from leaf axils, leaves on shrubs, and the 
leaf litter on the forest floor. The call has not yet been 
formally described, but consists of a pure, ringing, 
tonal note with no frequency or amplitude modula- 
tion; it sounds to the human ear like the sound pro- 
duced by the ringing of a small bell (1-2 notes/s). 

Localities and specimens: (Site 6) CAS 1 37641^42; 

(Site 11) PNM 2314-18, 2495, 2589-90; CMNH 
4113-15,4116-20,4868-69. 

Platymantis sp. 1 

Several immature specimens of a tiny, dark brown, 
tuberculate (1 1-15 mm SVL) species of Platymantis 
were taken at high elevations on Mt. Madja-as and on 
Mt. Baloy. They appear most similar to Platymantis 
pygmaeus of the Sierra Madre mountains of Luzon's 
east coast. Due to the fact that the available specimens 
are all sexually immature, we cannot recognize them 
taxonomically until further material and recordings of 
advertisements calls become available. 

Localities and specimens: (Site 11) CMNH 8132 
(Site 1 3) CMNH 3 1 73-74, 3177. 

Platymantis sp. 2 

Two immature specimens of a diminutive, black, 
smooth-skinned (12-13 mm SVL) species of Platy- 
mantis were collected at 1450 m from under leaf litter. 
The small sample size and immaturity of the speci- 
mens necessitates that taxonomic recognition of the 
species must await the collection of further material. 

Locality and specimens: (Site 1 1) CMNH 8133-34. 

Platymantis sp. 3 

This unidentified species is represented by a single, 
very large, black specimen with two dorsolateral light 
lines; it is appears possibly related to P. pseudodorsa- 
lis from Luzon (Brown et al., 1999). 

Locality and specimen: (Site 6) CAS 185495. 

Rana (= " Fejervarya") cancrivora cancrivora 
Gravenhorst 

This common frog is found in swamps, ponds, 
flooded rice fields and ditches (Inger, 1954). It is 
found in almost any pool of water at low elevations 
(Alcala and Brown, 1998). Inger (1954) first reported 
it on Panay Island. 

Localities and specimens: . (Site 1) PNM 1178-82; 
(Site 8) CAS 127893-95, 127899-904; (Site 9) PNM 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 45 



1156; (Site 16) CAS 127509, 127559-81, 127611, 
127678-79; (Site 17) CAS 127800; (Site 18) CAS 
127801-803; (Site 19) PNM 2607-16; (Site 22) 
USNM 78862-902; (Site 23) CAS-SU 14259-60, 
14452-79, 15486-514, USNM 77984-96, 77988- 
78042, 78048, 78062-65; (Site 24) FMNH 61398- 
402; (Site 26) PNM 1160-62; (Site 27) CAS-SU 
14513-25; (Site 28) CAS 124262-71, 185665-69; 
(Site 30) CAS-SU 14252-57; (Site 34) CAS-SU 
9762-63; (Site 36) CAS 124343; (Site 37) CAS 
124564-76; (Site 38) CAS 132878-79; (Site 39) CAS 
124135, 124175; (Site 40) USNM 339990, 340059- 
60; (Site 41) CAS 124178-79, 124320-28, 125020- 
28; (Site 42) CAS-SU 23966; (Site 44) CAS 125 194- 
96; (Site 46) CAS 125183-93; (Site 47) CAS 
125174-82; (Site 48) CAS 124791-92; (Site 50) 
USNM 78066-71. 

Rana(= " Hylarana") e/>tf/vae,3(Schlegel) (Fig. 
23) 

This widely-distributed and common frog is believed 
to have been introduced to the Philippines (Diesmos, 
1998; Alcala and Brown, 1998), originally on Negros 
(Inger, 1954; Alcala, 1962; Alcala and Brown, 1998). 
It's presence on Panay has been previously docu- 
mented (Taylor, 1920; Inger, 1954). We found our 
specimens along the grassy boarders of flooded rice 
fields. 

Localities and specimens: (Site 2) PNM 1725-26; 
(Site 3) CAS 137589; (Site 4) PNM 1603-10, 1612; 
(Site 7) PNM 1840; (Site 8) CAS 127891-92, 
127905-912; (Site 10) PNM 1174-76; (Site 11) 
CMNH 4870, PNM 1309-28, 2591-92; (Site 14) 
PNM 3816; (Site 19) PNM 2622-54; (Site 22) CAS- 
SU 9744; (Site 23) CAS-SU 14537-47, USNM 
77730-983, 78043-47, 78049-61; (Site 24) FMNH 
61391-97, 61449—52; (Site 27) CAS-SU 14578-84; 
(Site 30) CAS-SU 11120-24, 14526-36, FMNH 
40527; (Site 33) USNM 38650-54; (Site 38) CAS 
132881-86,' 134086-88; (Site 39) CAS 124209-212, 
124214; (Site 44) CAS 125158-69; (Site 47) CAS 
124143-57, 125310; (Site 50) USNM 77617-729, 
78413-38. 

Rana{- " Chalcorana") cf. et-weff/Boulenger 

Species of the Rana everetti complex are found in and 
along streams from 300 to about 1300 m (Inger, 1954; 
Brown and Alcala, 1955; Alcala, 1962; Alcala, 1986; 
Brown et al., 2000). Our Mt. Baloy expedition in 1989 
first recorded the presence of this frog on Panay 
(Sison et al., 1995). Negros populations (expected to 
be conspecific with those on Panay) are only found 
near water during breeding; this species has most 



often been collected in overhanging, streamside vege- 
tation (Brown and Alcala, 1955; Alcala, 1967). As 
noted by Brown et al. (2000), the taxonomic status of 
the Visayan populations referred to Rana everetti is in 
need of further study. 

Localities and specimens: . (Site 2) PNM 1732-33; 
(Site 14) PNM 3771, 3800-03, 3806-14, 3817-24, 
3896, 3913; (Site 39) CAS 124213, 124215-16. 

Rana (= " Fejervarya") vittigera Wiegmann 
(Fig. 24) 

This species occurs in open, agricultural areas near 
sources of water (ponds, flooded rice fields). The dis- 
tinctive, rapid honking call of this species can be 
heard in choruses of up to hundreds of individuals. 
This record appears to be the first published account 
of this species from Panay. 

Localities and specimens: . (Site 11) CMNH 4871 — 
72, PNM 2593-94; (Site 19) PNM 2596-606; (Site 
40) USNM 339991; (Site 41) CAS 124197; (Site 46) 
CAS 125360. 

Family Rhacophoridae 

Polypedates leucomystax (Gravenhorst) (Fig. 
25) 

This common tree frog is widely distributed in Panay 
(Inger, 1954; Alcala, 1986) in agricultural areas, for- 
est edges, and disturbed forests. It ranges from near 
sea level to 1000 m on Negros (Alcala. 1962) and was 
first documented on Panay by Inger ( 1954). Our spec- 
imens were collected in banana plantations and rice 
fields near the base of Mt. Madja-as and Mt. Baloy. 

Localities and specimens: . (Site 1) PNM 1107-09, 
1209-27; (Site 2) PNM 1729-30; (Site 6) CAS 
137599; (Site 7) PNM 1797-98; (Site 11) CMNH 
4997-98, PNM 2560-61; (Site 13) TNHC 56336, 
56338; (Site 14) PNM 3763, 3799; (Site 16) CAS 
127512-14, 127540-58, 127592-95, 127608-10, 
127647^19, 127657, 127670-76, 127680-97, 
127701; (Site 17) CAS 127721-26; (Site 18) CAS 
127841; (Site 24) FMNH 61485; (Site 25) PNM 
1107-09; (Site 28) CAS 124110-13, 124420, 
124422-25, 124581; (Site 29) USNM 339992-93; 
(Site 30) CAS-SU 11113-16, 14764, FMNH 40569, 
44263; (Site 37) CAS 124114, 125034-37; (Site 39) 
CAS 124158, 124192, 124204-07; (Site 40) USNM 
339994; (Site 41) CAS 124176, 125031-32; (Site 42) 
CAS-SU 23950-51, 23959-60, 23964-65; (Site 47) 
CAS 125342-43. 



Vol. 9, p. 46 



Asiatic Herpetological Research 



2001 



Reptilia 
Testudines 

Bataguridae 

Cuora amboinensis amboinensis (Daudin) 
(Fig. 26) 

This common species is found throughout the Philip- 
pines (Gaulke and Fritz. 1998) and on Panay (Gaulke, 
in press) in low elevation forests, agricultural areas, 
and near streams and swampy areas; it is generally 
considered nocturnal (Alcala, 1986). We have col- 
lected this species in stream-side habitats at low ele- 
vations on Panay. 

Localities and specimens: (Site 7) PNM 1888-89, 
1891-95; (Site 11) CMNH 5500, 5501, 5502; PNM 
1288, 5657-58; (Site 23) USNM 78103-04, 78746- 
49; (Site 41) CAS 153872; (Site 45) CAS 185507; 
(Site 50) USNM 78081-102. 

Squamata (Lizards) 

Agamidae 

Bronchocela cristatella (Kuhl) and B. marmo- 
rata (Gray) 

These arboreal lizards (Alcala, 1986) are found from 
lowland cultivated areas to lower midmontane pri- 
mary and secondary forests; they are most often 
encountered sleeping at night in stream-side vegeta- 
tion. No suitable taxonomic reference is available for 
Philippine Bronchocela and so the true identities of 
Panay specimens referred to B. cristatella and B. mar- 
morata are uncertain. There is little consensus regard- 
ing the identity of Bronchocela throughout the 
Visayas. Although specimens from Negros and Panay 
key out to earlier descriptions of both Bronchocela 
marmorata and B. cristatella (Taylor, 1922c; Alcala, 
1986), both "species" appear to be highly variable and 
diagnostic characters vary ontogenetically. We con- 
sider it unlikely that two independent lineages occur 
in sympatry on Panay and, at present, we hold in 
abeyance the identity of these populations until a thor- 
ough taxonomic revision is available. 

Localities and specimens: . (Site 6) CAS 137605; 
(Site 22) CAS-SU 10948; (Site 23) USNM 77133- 
38, 78105-107; (Site 41) CAS 124333; (Site 47) CAS 
125337-38. 



Draco spilopterus (Weigmann) (Figs 27, 28) 

This species is common at lower to mid-montane ele- 
vations and often is found in coconut groves and for- 
est edges (Alcala, 1986; McGuire and Alcala, 2000). 
Draco spilopterus is the only Draco species currently 
recognized from the Visayan and Luzon aggregate 
island complexes, despite biogeographic and morpho- 
logical evidence suggesting that Luzon and Visayan 
populations constitute independent evolutionary lin- 
eages (Taylor, 1922c; Heaney, 1985, 1986; McGuire 
and Alcala, 2000). Draco spilopterus may warrant 
further taxonomic attention once biochemical studies 
of species boundaries become available (Taylor, 
1922c; McGuire and Alcala, 2000). 

Localities and specimens: . (Site 3) CAS 137578, 
185504; (Site 4) PNM 1628-38; (Site 6) CAS 
137608-609, 185505; (Site 7) PNM 1759-79; (Site 8) 
CAS 127886, 127916, 127961, 128031; (Site 11) 
PNM 1275-82, 2720-21; (Site 14) PNM 3769-70, 
3878; (Site 18) CAS 127851-52; (Site 19) TNHC 
58465-67, 58471-80, 58482-90, 58850; (Site 43) 
CAS 39686; (Site 44) CAS 125295, USNM 38990- 
96; (Site 47) CAS 125277-94. 

Hydrosaurus pustulatus (Eschscholtz) (Figs. 
29, 30) 

The Mt. Madja-as specimens were collected in over- 
hanging stream-side vegetation. On Mt. Baloy we also 
collected a specimen in similar riparian habitats. 
These large omnivorous agamids can be found on 
trees and shrub-layer vegetation, overhanging streams 
and rivers (Alcala, 1986). The taxonomy of Philippine 
Hydrosaurus is in need further taxonomic studies. 

Localities and specimens: (Site 11) CMNH 5043; 
(Site 13) TNHC 56762; (Site 22) USNM 77091-103, 
85073-74; (Site 44) CAS 125336, USNM 38988-89; 
(Site 50) 77104-28, 78168-87. 

Gonocephalus sp. 

This genus was reported as an island record for Panay 
from site 2 by Sison, et al. (1995). The name G. 
sophiae has been applied to Negros populations (Tay- 
lor, 1922c; Alcala, 1986) and we might expect that if 
Negros populations are indeed G. sophiae (and this 
name applies to a distinct lineage that is independent 
from G. semperi), specimens from Panay might be 
referable to this species as well (see Gaulke, in press). 
At present, no suitable taxonomic reference exists, 
and species boundaries are unclear. Due to this fact, 
we do not apply a specific epithet to this population. 
Philippine populations of the genus Gonocephalus are 
greatly in need of taxonomic review. 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 47 



Localities and specimens: (Site 2) PNM 1130-32; 
(Site 14) PNM 3858. 

Gekkonidae 

Cosymbotus platyurus (Schneider) 

We found specimens of this common house gecko 
species in both the city and in agricultural areas; they 
are widely distributed on Panay (Brown and Alcala, 
1978) and are usually encountered under lights. 

Localities and specimens: (Site 7) PNM 1803, 
1815-16; (Site 8) CAS 128020; (Site 11) CMNH 
5137, PNM 1261-74; (Site 19) CMNH 5089-96, 
5098-104, PNM 2722-36; (Site 22) USNM 78776- 
832, 103480; (Site 23) USNM 77144-57, 339998- 
340010; CAS-SU 9613; (Site 26) PNM 1234-14, 
1247, 1249; (Site 29) USNM 339995-97; (Site 34) 
CAS-SU 9612, 12021; (Site 40) USNM 34011-12; 
(Site 50) FMNH 41302. 

Cyrtodactylus annulatus (Taylor) (Fig. 32) 

Brown and Alcala (1978) and Alcala (1986) report 
this species in forested areas ranging from sea level to 
1200 m on the nearby land-bridge islands of Negros, 
Cebu, and Inampulugan (Site 42) and its presence on 
Panay is not surprising. Nevertheless, this report and 
that of Gaulke (in press) appear to be the first records 
of this species from Panay. Brown and Alcala (1978) 
reported that this species has been collected on the 
forest floor, from under logs, beneath bark, and on the 
trunks of trees. We found Panay specimens in second- 
ary forest on root masses of trees protruding through 
eroding banks along a large river at the base of Mt. 
Madja-as. Predation on this species by flying snakes 
(Chrysopelea paradisi) was observed in these same 
microhabitats. 

Locality and specimens: (Site 28) CAS 124614-15, 
124768-78 (Site 42) CAS-SU 28009-10, 28013-14, 
28016-19, 28031, 28036-41, 28044-46,28050-60. 

Cyrtodactylus philippinicus (Steindachner) 
(Fig. 33) 

This species of Cyrtodactylus is found in a variety of 
habitats in the forest, particularly in rotting logs 
(Brown and Alcala, 1978; Alcala, 1986) and has been 
collected from sea level to nearly 1200 m (Brown and 
Alcala, 1978). In the Visayas, this species also has 
been collected on the nearby land-bridge islands of 
Negros, Pan de Azucar, and Boracay but never before 
on Panay. The specimens from Mt. Madja-as were 
collected in primary forest, during the day, under 
loose tree bark. 



Localities and specimens: .(Site 6) CAS 137607; 
USNM 496868; (Site 8) CAS 127883; (Site II) 
CMNH 5125-28, PNM 2751-53; (Site 13) TNHC 
56339; (Site 18) 127864, 127869; (Site 28) CAS 
124783-84; (Site 41) CAS 124046, 124780-82. 

Gehyra mutilata (Weigmann) (Fig. 34) 

This common and widely-distributed lizard is consis- 
tently found not only around human habitation (in 
darker areas, away from lights), but also in gardens 
and forested areas on trees (Brown and Alcala, 1978; 
Alcala, 1986). 

Localities and specimens: (Site 2) PNM 1682-86; 
(Site 3) CAS 137579; (Site 4) PNM 1649-53; (Site 7) 
PNM 1824; (Site 8) CAS 127888, 127922-29, 
127948-52, 127964, 127975-76, 127999-128000, 
128054-55; (Site 11) CMNH 5105-16, 5198, PNM 
1250-58, 2737-46; (Site 16) CAS 127504, 127607; 
(Site 18) CAS 127804, 127866; (Site 19) PNM 2737- 
46; (Site 23) USNM 77158, 78834-35; (Site 26) 
PNM 1245-49; (Site 28) CAS 124434-36, 124616- 
19; (Site 29) USNM 340013-14; (Site 37) CAS 
125029; (Site 39) CAS 124118-120, 124161-170; 
(Site 40) USNM 340015; (Site 41) CAS 124180-83; 
(Site 42) CAS-SU 28012, 28047-49; (Site 44) CAS 
124505-510; (Site 46) CAS 124687-88; (Site 47) 
CAS 124682-86, 125129-31; (Site 48) CAS 124511- 
12; (Site 50) FMNH 41383. 

Gekko gecko (Linnaeus) 

This common species is found around human habita- 
tion and in forest adjacent to disturbed areas. We col- 
lected specimens in secondary forest near the base of 
Mt. Madja-as. 

Localities and specimens: (Site 1) PNM 1056-63; 
(Site 2) PNM 2667; (Site 7) PNM 1062-1063, 1792- 
96; (Site 8) CAS 128004; (Site 11) CMNH 5018-25, 
PNM 1282, 1330, 2662-69; (Site 16) CAS 127582; 
(Site 17) CAS 127745; (Site 19) PNM 2665; (Site 23) 
CAS-SU 9585-88; USNM 77142-43, 340018-19; 
(Site 28) CAS 124979; (Site 29) USNM 340016-17; 
(Site 31) PNM 1 143; (Site 35) CAS 124393, 124318, 
124866-75; (Site 37) CAS 124315-17, 124929^19; 
(Site 39) CAS 124389-92; (Site 41) CAS 124319, 
124580; (Site 42) CAS-SU 27929; (Site 44) CAS 
125251; (Site 46) CAS 125249-50; (Site 47) CAS 
125247-48; (Site 49) CAS-SU 9589; (Site 50) 
FMNH 41377-81, 41376. 

Gekko gigante Brown and Alcala (Figs. 35, 
36) 

Gekko gigante was described by Brown and Alcala 
(1978) from the tiny land-bridge islands of Gigante 



Vol. 9, p. 48 



Asiatic Herpetological Research 



2001 



North and Gigante South, off the northeast coast of 
Panay (Fig. 2). This species has not been studied 
since its 1968 discovery until a recent visit to the type 
locality by RMB and A. Alcala in June 2000. We 
found G. gigante in small sea-side caves in karst lime- 
stone outcrops along the south coast of Gigante South. 
The northeast coast is the best place to survey for this 
species on Panay if suitable limestone habitat can be 
located. 

Localities and specimens: (Site 35) CAS 124318, 
124866-75 (Site 37) CAS 124315-17, 124929-49. 

Gekko mindorensisTay\or (Fig. 37) 

Brown and Alcala (1978), and Alcala (1986) report 
that individuals of this species are common on walls 
of caves, on tree trunks, and in leaves around the but- 
tresses of trees. Elsewhere in the Visayas this species 
has been reported from Negros, Cebu, and Caluya 
(Brown and Alcala, 1978). We have caught this spe- 
cies on cement walls, abandoned buildings in the for- 
est, in road-cut culverts, and on large dead tree trunks 
and stumps. Sison et al. (1995) reported this as an 
island record from Site 29. The taxonomic distinctive- 
ness of this species requires verification; recent data 
suggest the widespread G. mindorensis may be con- 
specific with G. kikuchii from Taiwan (Crombie and 
Ota, unpublished data; see also comment by Taylor, 
1922c). 

Localities and specimens: (Site 8) CAS 127882, 

127884-85, 128021; (Site 1 1) One uncataloged speci- 
men, deposited in PNM; (Site 16) CAS 127700; (Site 
18) CAS 127817; (Site 28) CAS 124767; (Site 29) 
PNM 2500; (Site 39) CAS 124136. 

Hemidactylus frenatus Schlegel in Dumeril 
and Bibron 

This common house gecko is widespread on Panay 
and often is associated with Cosymbotus platyurus 
(Alcala, 1986) in well lighted areas in human habita- 
tion. We collected several specimens near the base of 
Mt. Madja-as on walls of houses in agricultural areas. 

Localities and specimens: (Site 3) CAS 137580, 
USNM 496869-70; (Site 7) PNM 1801-02, 1804-14; 
(Site 8) CAS 127878-81, 127897, 127913-14, 
127930-45, 127953-58, 127963, 127977-78, 
128001-03, 128022-26, 128034-35, 128056-57, 
136742^14; (Site 11) CMNH 5148-54, PNM 1259- 
60, 1646-48, 2771-76; (Site 16) CAS 127505-07, 
127515-16, 127583, 127615-16, 127642^*6, 
127652-56, 127658-59, 127699; (Site 17) CAS 
127710-20, 127727-M, 127748-65, 127795-99, 
136741; (Site 18) CAS 127805-14, 127818-26, 



127829-10, 127848-50, 127865; (Site 19) CMNH 
5129-36, 5139^17, PNM 2754-70; (Site 23) USNM 
78833, 78836-39; (Site 26) PNM 1245-1246; (Site 
28) CAS 124081-92, 124427, 124663, 125002-12; 
(Site 29) USNM 340021-27; (Site 33) USNM 
340028-32; (Site 35) CAS 124363, 124504; (Site 37) 
124364-72, 124357-62, 124582-610, 124662; (Site 
39) CAS 124137^3, 124217^2; (Site 40) 340034- 
41; (Site 41) CAS 124373-85, 124664-81, 125013; 
(Site 42) CAS-SU 28007-08, 2801 1, 28021, 28034- 
35, 28042-43; (Site 44) CAS 124466, 124526-58 
[Nadulao Island]; (Site 46) CAS 124458-65, 125315- 
30, 124736-66; (Site 47) CAS 124448-57, 125128, 
125313-14; (Site 50) FMNH 41384-86. 

Hemidactylus ste/negeri 0\a and Hikida 

Previously referred to H. garnoti (Brown and Alcala, 
1978). the status of Philippine populations was clari- 
fied by Ota and Hikida (1989) and Ota et al. (1993). 
Sison et al. (1995) reported this species as an island 
record for Panay. 

Locality and specimen: (Site 25) CMNH 3225. 

Hemiphyllodactylus insu/arisJay\or 

While Brown and Alcala (1978) referred Philippine 
(and Panay Island) populations of Hemiphyllodacty- 
lus to H. typus, collections in USNM contain males 
and are not, therefore, the widespread unisexual H. 
typus, but probably are better referred to Taylor's H. 
insularis (R. Crombie, personal communication.). 

Localities and specimens: . (Site 3) CAS 137581- 
83; (Site 8) CAS 127889, 127965-71; (Site 18) CAS 
127855-57. 

Lepidodactylus lugubris (Dumeril and 
Bibron) 

This lowland gecko is common in patchy distributions 
and found in leaf axils, under bark, in tree holes, in 
coconut groves, and associated with large rocks in 
coastal areas (Brown and Alcala, 1978; Alcala, 1986). 
Though reported from Negros, Cebu, Inampulugan, 
Boracay, Gigante, and Pan de Azucar (Brown and 
Alcala, 1978), this species has not yet been reported 
from Panay. 

Localities and specimens: (Site 8) CAS 127947, 
127959, 127979, 127982-87, 127989, 128005-07, 
128010, 128012-14, 128017, 128043-44, 128046- 
50, 128059, 154716; (Site 28) CAS 124579; (Site 37) 
CAS 124200, 125030, 125033; (Site 39) CAS 
1241 16-17; (Site 41) CAS 124981-125000; (Site 42) 
CAS-SU 28015, 28061; (Site 44) CAS 124561-63; 
(Site 46) CAS 124492. 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 49 



Lepidodactylus planicaudus Stej neger 

Alcala and Brown (1978) recorded this species in 
coconut trees, mangroves, and in fern axils; they have 
also been taken on tree trunks in forests from sea level 
to 700 m. Elsewhere in the Visayan Aggregate Island 
Complex it is known from Cebu, Guimaras, Caluya, 
Masbate and Boracay (Brown and Alcala. 1978). 

Localities and specimens: (Site 3) CAS 139939; 
(Site 8) CAS 127921, 127980-81, 127988, 127990, 
128008-09, 128011, 128015-16, 128018-19, 
128045, 128051-53; (Site 16) CAS 127698; (Site 17) 
CAS 127709; (Site 39) CAS 124115, 124134; (Site 
46) CAS 124823-26. 

Scincidae 

Brachymeles boulengeri taylon 'Brown (Figs. 
38, 39) 

Usually encountered in agricultural areas (especially 
coconut plantations) adjacent to forest, this species 
most often is collected from its preferred microhabi- 
tat, inside rotting coconut logs. It has also been col- 
lected in mature and secondary forest, from sea level 
to 1200 m on numerous islands in the Visayas (Brown 
and Alcala, 1980) but this is the first record from 
Panay. 

Localities and specimens: (Site 1) PNM 1 148; (Site 
25) PNM 1 148; (Site 39) CAS 124157; (Site 41) CAS 
124044 (Site 42) CAS 27930-31, 27946-51, 27953. 
27973-84, 27987-93. 

Brachymeles talinis Brown 

This skink occupies decaying vegetation and humus 
on the floor of primary and, occasionally, secondary 
forest (Brown and Alcala, 1980; Alcala, 1986). Alcala 
and Brown (1980) reported its elevational range from 
sea level to approximately 1200 m. 

Localities and specimens: (Site 6) 137603; (Site 8) 
CAS 127870-75, 127962, 128029-30, 154689; (Site 
14) PNM 3852, 3856, 3859, 3909-10; (Site 16) CAS 
127517-39, 127584-90, 127606, 127663-68; (Site 
28) 154692, 200522-31; (Site 39) CAS 124148, 
200521; (Site 42) CAS-SU 27972, 27996-97; (Site 
50) CAS 137603. 

Brachymeles tridactylus Brown (Fig. 40) 

This species has been collected under logs, humus, 
and other debris in forested regions from sea level to 
approximately 900 m (Brown and Alcala, 1980; 
Alcala, 1986). On Mt. Madja-as we found them under 
logs in well-regenerated second growth forest below 
200 m. 



Localities and specimens: (Site 3) CAS 137566-75; 
(Site 6) CAS 137602, 137604; (Site 8) CAS 127876- 
77, 127915, 127974, 128027-28, 128037^12; (Site 
11) PNM 5514-15; (Site 42) CAS-SU 27950, 27952, 
27985-86, 28456. 

Das/a grisea (Gray) 

In the Philippines, this species has been recorded from 
the islands of Mindoro, Luzon, and Marinduque 
(Brown and Alcala, 1980). Its inclusion in this paper 
is based on a single record from Semirara Island, a 
small island south of Mindoro that is included in the 
political boundary of Antique province but is on the 
land-bridge to Mindoro. We do not expect this species 
to be discovered on Panay and we do not regard it as a 
Visayan Aggregate Island complex resident. 

Locality and specimen: (Site 18) CAS 134218. 

Dasia semicincta (Peters) 

A large lowland species, previously reported only 
from Mindanao in the Philippines (Brown and Alcala, 
1980; Alcala. 1986). Outside of the Philippines D. 
semicincta is known from Borneo (Brown and Alcala, 
1980). This is the first record of this species from 
Panay; it also is a substantial range extension in need 
of verification. 

Locality and specimen: (Site 23) USNM 78840. 

Emoia a trocostata (Lesson) 

This species of skink is common in mangroves and 
can be found active on tree trunks, in tree holes, and in 
rock crevices (Alcala and Brown, 1967; Brown and 
Alcala, 1980; Alcala, 1986). Taylor (1922c) reported 
that this species swims in brackish water, burrows in 
sand, and feeds on small crabs. On Panay and Negros, 
mangrove forests have nearly all been cleared, possi- 
bly rendering this species rare in recent collections. 

Localities and specimens: (Site 16) CAS 127638- 
39; (Site 18) CAS 127842-45; (Site 22) CAS-SU 
13585; (Site 23) USNM 77139, 7841 1, 78841, 80934; 
(Site 37) CAS 125040; (Site 41) CAS 124329-32, 
124620-24; (Site 42) CAS-SU 27920; (Site 47) CAS 
125335; (Site 50) USNM 77141. 



smaragdina philippinica 



Lamprolepis 
(Mertens) 

This skink is common in coconut groves and gardens 
and a variety of natural vegetation types (Alcala, 
1986). Near Mt. Madja-as and Mt. Baloy we observed 
them on coconut trunks at low elevations. 

Localities and specimens: (Site 4) PNM 1639; (Site 
7) PNM 1780. 1782, 1786; (Site 8) CAS 127887, 



Vol. 9, p. 50 



Asiatic Herpetological Research 



2001 



127917-20, 127946, 127991-98, 128058, 128060- 
61; (Site 11) CMNH 5038^0, PNM 1283, 2681-82; 
(Site 14) PNM 3798; (Site 16) CAS 127508, 127612- 
14, 127640-41, 127669; (Site 17) CAS 127747, 
127794; (Site 18) CAS 127853-54; (Site 19) TNHC 
56472-73; (Site 28) 124053-57; (Site 35) CAS 
124198, 12471 1-13; (Site 39) CAS 124201-02; (Site 
40) USNM 340061; (Site 42) CAS-SU 27954-62, 
27969; (Site 44) CAS 125301; (Site 46) 125298-300; 
(Site 47) CAS 124447, 125296-97. 

Lipinia pulchella taylon '(Brown and Alcala) 

This widely distributed arboreal skink had not been 
reported from Panay prior to our visits to Mt. Majda- 
as. Previously, L. p. taylori had only been known from 
Negros (Brown and Alcala, 1980). Brown et al. (1996; 
2000) have collected specimens from Luzon that key 
out to this subspecies, potentially indicating the need 
for a review of Brown and Alcala's (1980) taxonomy 
of this species. This is a new island record for Panay. 

Localities and specimens: (Site 11) CMNH 5083; 
(Site 25) PNM 1156. 

Mabuya indeprensa Brown and Alcala 

Sison et al. (1995) reported this species as an island 
record, collected at about 200 m on Mt. Baloy. Well 
known from forested, montane habitats on Negros and 
Cebu (Brown and Alcala, 1980), this species is found 
in leaf litter, stumps and fallen logs, and around tree 
buttresses. 

Localities and specimens: (Site 1)PNM 1149; (Site 
9) PNM 1 153-55; (Site 11) CMNH 51 19-22, 5199- 
202, PNM 2747-49, 5511-13; (Site 13) one uncata- 
loged specimen deposited in PNM; (Site 16) CAS 
127596-97, 127599, 127601-02, 127604-05; (Site 
18) CAS 127863; (Site 25) CMNH 3247, PNM 1 151- 
1152. 

Mabuya multicarinata borealis Brown and 
Alcala (Fig. 41) 

This subspecies commonly is found under tree bark, 
logs and piles of vegetation (Brown and Alcala, 1980; 
Alcala, 1986); in the Visayas it has been collected on 
Negros, Caluya, Semirara, Gigante, Pan de Azucar, 
and Cebu, but this is the first record from Panay. 
Brown and Alcala (1980) reported finding one speci- 
men as high as 1500 m on Luzon island. 

Localities and specimens: . (Site 2) PNM 1657-60, 
1668-80; (Site 6) USNM 496871; (Site 16) CAS 
127598, 127600, 127603, 127650-51; (Site 18) CAS 
127862. 127867-68; (Site 28) CAS 124107-109, 
124971-78; (Site 35) CAS 124493-97; (Site 36) CAS 



124199, 125039; (Site 37) CAS 125038; (Site 41) 
CAS 124050. 

Mabuya multifasciata (Kuhl) 

This species was found in a variety of habitats from 
beaches and low elevation agricultural areas, to dis- 
turbed forest adjacent to primary forest on Mt. Madja- 
as. Though known to occur throughout the Philip- 
pines, in the Visayas, this species has only been 
reported from Negros (Brown and Alcala, 1980); this 
is the first published account from Panay. 

Localities and specimens: (Site 2) PNM 1661-65; 
(Site 3) CAS 137585, USNM 496872; (Site 4) PNM 
1640-45, 1655-56; (Site 6) CAS 137610-12, 139147; 
(Site 7) 1781, 1783; (Site 1 1 ) CMNH 5203-04, PNM 
5510; (Site 14) PNM 3845-46, 3853; (Site 22) CAS- 
SU 13632; (Site 25) PNM 1149-55; (Site 42) CAS- 
SU 27968, 27971, 27998; (Site 44) CAS 125264-66; 
(Site 46) CAS 125334; (Site 47) CAS 125262-63, 
125333; (Site 50) FMNH 4 1 389^104. 

Parvoscincus s/son/Femer, Brown and Greer 

All specimens of this recently-described species were 
collected during the day from beneath loose soil and 
leaf litter in the forest between 900 and 1125 m 
(Ferner et al., 1997). At present, this species is known 
only from Mt. Madja-as; the only other member of the 
genus is from Palawan Island (P. palawanensis). 

Locality and specimens: (Site 11) CMNH 3797-99, 
PNM 2308-10. 

Sphenomorphus arbore/?sTay\or 

This forest species is found under logs and leaves and 
occasionally on buttresses at the base of tree trunks 
(Brown and Alcala, 1980; Alcala, 1986). Our speci- 
mens were collected in primary forest between 1400 
and 1600 m in pit-fall traps. 

Localities and specimens: . (Site 6) CAS 137644; 

(Site 1 1) CMNH 5062-74, PNM 2684-710; (Site 14) 
PNM 3720, 3746-48, 3753, 3867-69, 3871, 3875, 
3898, 3901-03; (Site 41) CAS 124048-49. 

Sphenomorphus coxi divergens~\ay\ox 

This species is often found at the forest edge and in 
forest gaps. It is widely-distributed in the archipelago 
(Brown and Alcala, 1980; Alcala, 1986). Sphenomor- 
phus coxi divergens previously has been reported 
from Luzon, Marinduque, and Mindoro but not the 
Visayas (Brown and Alcala, 1980). This is the first 
published record of a specimen from Panay. 

Locality and specimens: (Site 11) CMNH 5123, 
PNM 2750. 



2()()l 



Asiatic Herpetological Research 



Vol. 9, p. 51 



Sphenomorphus cumingi '(Gray) 

This large forest species is found around fallen logs 
and tree buttresses and is believed to be limited to low 
to medium elevations (Brown and Alcala, 1980; 
Alcala, 1986). Sphenomorphus cumingi previously 
has been reported from islands in the Luzon and Min- 
danao aggregate island complexes; this is the first 
published record of a specimen from Panay. 

Locality and specimen: (Site 28) CAS 124779. 

Sphenomorphus jagori grandis Taylor (Fig. 
42) 

This large skink is found under rotting logs, in sun 
spots in forested and disturbed areas, and along 
streams (Brown and Alcala, 1980; Alcala, 1986). 

Localities and specimens: .(Site 6) 137645^8; 

(Site 14) PNM 3724-5, 3755; (Site 16) CAS 127660- 
62, 127677; (Site 28) CAS 125267-76, 154659-68; 
(Site 37) CAS 124835, 185508; (Site 41) CAS 
185509-516. 

Sphenomorphus 5/eere/Stejneger 

We found this species on Mt. Madja-as in leaf litter in 
montane forest. Brown and Alcala (1980) report that 
this species is common in leaf litter of primary forest 
and in secondary growth. 

Localities and specimens: (Site 3) CAS 137600- 
601, 139169-70, USNM 496873-74; (Site 11) 
CMNH 5026-37, PNM 2670-80; (Site 14) PNM 
3708-09, 3721-23, 3726, 3749-52, 3754, 3775-81, 
3786-87, 3797, 3826-29, 3836-42, 3847-51, 3854- 
55, 3861, 3870, 3872-74, 3876-77, 3879-81, 3890- 
92, 3914; (Site 25) PNM 1159; (Site 28) CAS 
124437-38, 124577-78; (Site 36) CAS 124830; (Site 
37) CAS 124827-29, 124839; (Site 39) CAS 125091; 
(Site 47) CAS 124625-61, 125064, 125090. 

Tropidophorus grayi Giinther (Fig. 43) 

Sison et al.'s (1995) Mt. Baloy specimen was the first 
record of this species from Panay. We also found a 
specimen at the base of Mt. Madja-as underneath a 
rock on the edge of a small stream in second growth 
forest. 

Localities and specimens: (Site 11) CMNH 5117; 
(Site 13) One uncataloged specimen deposited in 
PNM. 



Varanidae 

Varanus salvator nuchalis (Giinther) (Figs. 44, 
45) 

This monitor lizard is found in agricultural and dis- 
turbed areas, mangrove swamps, and forested areas 
from sea level to about 1200 m (Alcala, 1986; Gaulke, 
1991a, 1991b, 1992). On Mt. Madja-as we collected 
one specimen during the day in second growth forest 
where it was active near a small stream. 

Localities and specimens: (Site 1 ) PNM 1 142; (Site 
11) PNM 5660; (Site 22) CAS 1 1018; (Site 31) PNM 
1142; (Site 37) CAS 124879; (Site 44) CAS 124881; 
(Site 47) CAS 124880; (Site 50) USNM 77129-32, 
80115-19; FMNH 41417-18. 

Squamata (Snakes) 

Acrochordidae 

Acrochordus granulatus (Schneider) 

Found in mangroves and at river mouths, this species 
feeds exclusively on fish and is widely distributed and 
common (Alcala, 1986). We are not aware of other 
published accounts of this species from Panay. 

Locality and specimens: (Site 23) USNM 78412, 
78744-45, 78906, CAS-SU 8695, 8769. 

Boidae 

Python reticulatus (Schneider) (Fig. 46) 

While this snake is considered widely distributed and 
common in the tropical rain forests as well as near 
human habitation (Alcala, 1986), only one specimen 
has been collected on Panay Island (Leviton, 1963c; 
see also Gaulke, in press). 

Locality and specimen: (Site 44) CAS 124916. 

Colubridae 

Ahaetulla prasina preocularis (Taylor) (Fig. 
47) 

This snake may be found in low trees and shrubs from 
sea level to about 800 m on Panay (Leviton, 1963c, 
1968; Alcala, 1986). The specimens from Mt. Madja- 
as were collected from vegetation overhanging a 
small stream at low elevation. 

Localities and specimens: (Site 2) PNM 1689; (Site 
7) PNM 1787; (Site 1 1) CMNH 5084-85, PNM 2716; 
(Site 14) PNM 3825; (Site 29) USNM 340042; (Site 
44) CAS 125339; (Site 50) FMNH 41 108. 



Vol. 9, p. 52 



Asiatic Herpetological Research 



2001 



Boiga angulata (Peters) 

We found our specimen in a coconut tree in a clearing 
adjacent to virgin forest at 900 m on Mt. Madja-as. 
While this species is known from Negros (Leviton, 
1970a), until this report and that of Gaulke (in press) 
it has never been recorded on Panay. 

Locality and specimen: (Site 1 1) CMNH 5504 

Boiga xf cynodon (Cuvier inV. Boie) 

This species has been found in forested areas at low 
altitudes on Palawan, Mindanao, and Luzon (Leviton, 
1963c, 1970a; Alcala, 1986) but previously it has not 
been reported in the Visayas (see also Gaulke, in 
press). 

Locality and specimen: (Site 41) CAS 125173. 

Boiga ci. dendrophila (Boie) 

Mangrove snakes usually are found in branches of 
low trees and bushes in forested areas (Leviton 1968; 
Alcala, 1986) at low elevations. This is the first record 
of a species in this complex from Panay. 

Localities and specimens: (Site 28) CAS 124388; 
(Site 39) CAS 124386-87. 

Calamaria geravaisi DumerW and Bibron 

A burrowing snake, this species is commonly found in 
the humus under rotting logs and feeds on earthworms 
(Leviton, 1963c; Inger and Marx, 1965; Alcala, 1986). 
On Mt. Madja-as we found specimens under rotten 
logs and large flat rocks. 

Localities and specimens: (Site 2) PNM 1087; (Site 
11) CMNH 5081-82, PNM 2714-15; (Site 23) CAS- 
SU 15953-57, 15962-65; (Site 44) CAS 124612. 

Cerberus rynchops (Schneider) 

This aquatic snake has been collected in brackish 
swamps, mangroves, fish ponds, and river estuaries in 
coastal areas (Gyi, 1970; Alcala, 1986) 

Localities and specimens: (Site 1) PNM 1053-55, 
1077-83; (Site 20) CAS-SU 13079; (Site 22) CAS- 
SU 8696-97, 8719; CAS-SU 12380, CM R2423, 
R2426; (Site 23) USNM 77159-178, 78907-19; (Site 
40) 340043; (Site 50) FMNH 41115-17. 

Chrysopelea paradis/'Boie 

This species was not known from Panay (Leviton, 
1963, 1964a) until Sison, et al. (1995) reported it as 
an island record from Site 1. On Mt. Madja-as we 
found this arboreal snake during the day in root 
masses overhanging a river bank; one specimen was 
preying on a Cyrtodactylus annulatus when captured. 



We have also observed this species in coconut groves 
at sea level near the town of San Jose (site 19). 

Localities and specimens: (Site DPNM 1050; (Site 
8) CAS 128032; (Site 11) CMNH 5041-5042, PNM 
2683; (Site 19 TMM 56474; (Site 27) CAS 185-672; 
(Site 36) CAS 125172; (Site 46) CAS 125331-32. 

Cyclocorus lineatus alcalai Leviton 

We found our specimens in habitats ranging from dis- 
turbed second growth at sea level to first growth for- 
ested riparian sites at 1400 m. At the time of its 
description (Leviton, 1967), this subspecies was 
known only from the nearby islands of Negros and 
Cebu (Fig. 2). This is the first published account of 
this species from Panay. 

Localities and specimens: (Site 1) PNM 1047, 
1065; (Site 2) PNM 1065, 1688; (Site 3) CAS 
137576; (Site 6) CAS 137606; (Site 11) CMNH 
5086-87, PNM 2717-18; (Site 14) 3884-85, 3912; 
(Site 16) CAS 127702, 127706; (Site 28) CAS 
124051-52, 124421; (Site 37) CAS 125171; (Site 47) 
CAS 124445. 

Dendrelaphis caudolineatus terrificus 
(Peters) 

This subspecies is known from Panay and Negros and 
usually is found in forested and cultivated areas in or 
along swamps and streams (Leviton, 1970b; Alcala, 
1986). Sison et al. (1995) reported this as an island 
record from 200 m on Mt. Baloy. The specimen from 
Mt. Madja-as was sleeping in bushes near a river 
when captured. 

Localities and specimens: (Site 4) CAS 1654; (Site 
8) CAS 127896, 128033; (Site 1 1) CMNH 5080; (Site 
13) CMNH 3254; (Site 16) CAS 127703-705, 
127707-708; (Site 18) CAS 127828, 127847, 127859; 
(Site 28) CAS 125170; (Site 39) CAS 124203, 
185673-74; (Site 42) CAS-SU 28004; (Site 50) 
FMNH 41093-96. 

Dendrelaphis pictus pictus (Gmelin) (Fig. 48) 

This common snake is arboreal and usually found 
near streams, in vegetation surrounding flooded rice 
fields, and in swampy areas (Leviton, 1963c, 1970b; 
Alcala, 1986). The Mt. Madja-as specimens were col- 
lected at night where they slept in stream-side vegeta- 
tion. 

Localities and specimens: (Site 1) PNM 1049, 
1071; (Site 3) CAS 137577; (Site 7) PNM 1788-90; 
(Site 9) PNM 1071; (Site 11) CMNH 5078-79, PNM 
2713; (Site 14) PNM 3843-14; (Site 22) CAS-SU 
8660-70, 8718, 14932, 14936-37, CMNH 2408-13, 



200! 



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Vol. 9, p. 53 



CM R2226, R2408-13; (Site 23) CAS-SU 8698-99, 
8708-710; USNM 77419-591, 340044-51; (Site 30) 
CAS-SU 14931 ; (Site 38) CAS 200256; (Site 44) 
CAS 125255-61; (Site 46) CAS 125254; (Site 47) 
CAS 125252-53; (Site 48) CAS 124725; (Site 50) 
FMNH 4109-104, 41106, USNM 77592-609. 

Elaphe erythrura psephenoura Leviton 

This is a common lowland snake, often found near 
human habitations (Leviton, 1979; Alcala, 1986). On 
Mt. Madja-as we collected one specimen from the for- 
est floor where it was active at midday. 

Localities and specimens: (Site 1) PNM 1048, 
1051; (Site 1 1) PNM 5662; (Site 22) CAS-SU 12389, 
13212-13; (Site 23) USNM 340052; (Site 30) CAS- 
SU 13217; (Site 38) CAS 131700; (Site 42) CAS-SU 
28001; (Site 47) CAS 125141-^12, 125340. 

Gonyosoma oxycephala (Reinwardt in F. 
Boie) 

This is an arboreal snake that is found in disturbed 
and primary forests (Alcala, 1986). On Mt. Madja-as 
we collected one specimen that was active at mid day 
in a tree above a large river (4 m from the ground). 
This record, and that of Gaulke (in press) are the first 
published accounts of this species from Panay. 

Localities and specimens: (Site 1)PNM 1084; (Site 
9) PNM 1084; (Site 1 1) CMNH 5503; PNM 5659. 

Hologerrhum derma// Brown, Leviton, Ferner, 
and Sison 

We first collected this newly-described species 
(Brown et al., this issue) between 1030 and 1510 m in 
climax forest on Mt. Madja-as. Specimens were col- 
lected in a dry stream bed and in leaf litter on the for- 
est floor 30 m from a large river. One specimen from 
the Municipality of San Remegio was collected at 
approximately 700 m above sea level. Recent survey 
work in NW Panay suggests that this species also 
occurs in forested areas at lower elevations (Gaulke, 
in press). 

Localities and specimens: (Site 1 1 ) CMNH 5075, 
PNM 2711 (14) PNM 3704. 

Lycodon aulicus capucinus (H. Boie in F. 
Boie) (Fig. 49) 

This common, widespread, nocturnal snake is often 
found in gardens, agricultural areas, and around 
houses (Leviton, 1965; Alcala, 1986). Our specimens 
were collected near rice fields away from forested 
areas. 



Localities and specimens: (Site 1) PNM 1072-75; 
(Site 3) CAS 137584; (Site 8) CAS 127960; (Site 12) 
PNM 1072; (Site 13) PNM 1380; (Site 22) CAS-SU 
8671, 8700-01, CMNH 2443, CM R2443; (Site 23) 
USNM 77616, 340053; (Site 26) PNM 1073-75; (Site 
47) CAS 125341. 

Oligodon modes turn (G un ther ) 

This species is found under rotting logs and forest 
floor debris; it is known from sea level to 400 m 
(Alcala, 1986). In the Visayas, this species was only 
known from Negros (Leviton, 1963a) before Sison et 
al. (1995) first reported it on Panay. 

Localities and specimens: (Site 1) PNM 1066; (Site 
2) PNM 1067; (Site 14) PNM 3790, 3866. 

Psammodynastes pu/veruientus(H. Boie in?. 
Boie) 

Leviton (1963c; 1983) and Alcala (1986) report this 
snake as common up to elevations of 1000 m or more 
in moist forests on Negros Island; Leviton's (1963c) 
listing of this species from Panay was not repeated in 
his 1983 review of the genus in the Philippines and no 
specimens were reported from Panay in the later 
paper (Leviton, 1983). This account and that of 
Gaulke (in press) appears to be the first vouchered 
records of this species from Panay. 

Localities and specimens: (Site 1) PNM 1068; (Site 
2) PNM 1069; (Site 25) PNM 1067, 1070. 

Pseudorabdion /ncnan?araeTay\or 

Prior to Sison et al. (1995) this species was known 
only from Negros and Luzon (Brown and Leviton, 
1959; Alcala, 1986). Specimens from Mt. Baloy were 
collected at 950 m under rotting logs in original for- 
est. 

Locality and specimens: (Site 13) Two uncataloged 
specimens in PNM (PNM Field Numbers 163 and 
209). 

Pseudorabdion oxycephalum (Gunther) 

Previously considered a rare snake endemic to Negros 
Island, this species is now known from other localities 
in the Luzon and Mindanao aggregate island com- 
plexes (Brown and Leviton, 1959; Leviton, 1963c; 
Alcala, 1986; Brown et al., 1999). This is the first 
published record from Panay. Pseudorabdion 
oxycephalum is been found in humus and under rot- 
ting logs from sea level to about 750 m. 

Localities and specimens: (Site 6) CAS 137643; 
(Site 39) CAS 124174, 124193; (Site 41) CAS 
124043. 



Vol. 9, p. 54 



Asiatic Herpetological Research 



2001 



Pseudorabdion talonuran Brown, Leviton and 
Sison 

The discovery of this new species at high elevations 
was surprising. Both specimens were found under 
logs on Mt. Madja-as in forest classified as the transi- 
tion zone between mixed dipterocarp (submontane) 
and mossy (upper montane; Whitmore, 1984). The 
holotype was found at 1500 m and the paratype at 
1410 m. 

Locality and specimens: (Site 1 1 ) CMNH 5076, 
PNM2712. 

Tropidonophis negrosensis (Taylor) 

This species of water snake is common along forest 
streams from sea level to about 700 m (Leviton, 
1963c; Alcala, 1986; Malnate and Underwood, 1988). 
The specimen from Mt. Madja-as was found on a 
river bank at midday. 

Localities and specimens: . (Site 3) CAS 185749; 
(Site 6) CAS 137613; (Site 11) CMNH 5124; (Site 
14) PNM 3911; (Site 22) CAS-SU 15971; (Site 28) 
CAS 12461 1; (Site 41) CAS 124047. 

Zaocys luzonensis Gunther 

A common tropical forest snake, this species ranges 
from sea level to over 1 100 m (Leviton, 1983; Alcala, 
1986; Ross et al., 1987). On Mt. Madja-as, we found 
two specimens active at midday in disturbed forest at 
low elevation. 

Localities and specimens: (Site 1)PNM 1052; (Site 
2) USNM 269078; (Site 7) PNM 1791; (Site 11) 
CMNH 5505; PNM 5663. 

Elapidae 

Calliophis calligaster gemianulis (Peters) 

We collected one specimen on Mt. Madja-as at 
approximately 800 m on a mountain path away from 
water. They have previously have been found in a 
variety of semifossorial habitats associated with 
regenerated and climax forest (Leviton 1963b, 1963c; 
Alcala, 1986) 

Localities and specimens: (Site 11) CMNH 5088, 
PNM 2719; (Site 13) PNM 1379; (Site 21) UPLB 
2184; (Site 22) CM R2581; (Site 23) CAS-SU 12966- 
68; (Site 44) CAS 125364; (Site 47) CAS 125363. 

Hydrophis belcheri Gray 

This species has been recorded from the Visayan sea 
and is thought to be rare (Alcala, 1986). 



Localities and specimens: (Site 5) USNM 38588; 
(Site 37) FMNH 202832-36, 202838, 202840-42. 

Hydrophis cyanocinctus Daudin 

Alcala (1986) mentioned records from the Visayan 
Sea. 

Locality and specimens: (Site 37) FMNH 202843- 
54. 

Hydrophis elegans Gray 

Locality and specimen: (Site 5) USNM 38589. 

Hydrophis inornatus Gray 

Locality and specimen: (Site 23) CAS-SU 8778. 

Lapemis hardwickii Gray 

Alcala (1986) reported this species from the Visayan 
sea; he noted that specimens were collected by trawl- 
ing. 

Localities and specimens: (Site 23) USNM 77610- 
15. 

Laticauda colubrina Schneider 

This sea snake is found among rocks and in coral reef 
areas near shore; it is commercially used for its meat 
and hide (Alcala, 1986). 

Locality and specimens: (Site 37) FMNH 202797- 
801,202804-808,202810. 

Typhlopidae 

Ramphotyphlops braminus (Daudin) 

This species is common under rocks and other debris 
in agricultural areas. It can also be found in similar 
microhabitats along the edges of forested areas 
(Alcala, 1986). Taylor (1922d) reported catching 
many specimens under rocks immediately following 
rains. 

Localities and specimens: . (Site 1) PNM 1044-46; 
(Site 8) CAS 127972, 128036; (Site 17) CAS 127746; 
(Site 18) CAS 127846, 127858, 127860; (Site 30) 
CAS-SU 12544-49; (Site 37) CAS 125041; (Site 46) 
CAS 124503. 

Ramphotyphlops cumingii (Gray) 

Alcala (1986) reports this blind snake is found in epi- 
phytic ferns in the trees of moist forests at low to 
medium elevations. In all of his field work, Taylor 
(1922d) was unable to capture this species, suggesting 
he may have overlooked its preferred microhabitat. 



2001 



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Vol. 9, p. 55 



Localities and specimens: (Site 28) CAS 169877; 
(Site 35) CAS 125092. 

Typhlops castanotusWynn and Leviton 

Described very recently (Wynn and Leviton. 1993). 
this distinctive bicolored species is known from 
Negros. Panay (Makato). Boracay. and Inampulugan 
islands. It has been collected under debris in bamboo 
and coconut groves, hardwood forests and forest rem- 
nants. 

Localities and specimens: (Site 3) CAS 139171; 
(Site 8) CAS 127973; (Site 42) CAS-SU 27934-16. 

Typhlops hypogius Savage, Typhlops luzon- 
ms/s Taylor, and Typhlops ruberBoettger 

Found in detritus under rotting logs, these snakes have 
been collected in forests and disturbed areas near for- 
est from sea level to about 800 m (Alcala. 1986). 
Wynn and Leviton (1993) followed McDowell ( 1974) 
in referring Philippine T. luzonensis and T. hypogius 
to T. ruber. Recently, McDiarmid et al. (1999) 
asserted that T. luzonensis and T. hypogius should be 
recognized until more compelling evidence is pre- 
sented that would suggest they are conspecific with T. 
ruber. We agree, noting that the type localities for T. 
hypogius (Cebu), T. ruber (Samar), and T. luzonensis 
(Luzon) are each located on separate Pleistocene 
aggregate island complexes that might be expected to 
support independent lineages of blind snakes. If this is 
so, and the Visayan islands contain a single distinct 
lineage in this species complex, the name Typhlops 
hypogius would most likely apply to specimens from 
Cebu. Negros. Panay. Masbate. and smaller, land- 
bridge islands. 

Localities and specimens: (Site 18) CAS 127861. 

Family Viperidae 

Tropidolaemus cf wagleri (H. Boie in F. Boie) 
(Figs. 51, 52) 

This arboreal viper is common in forest bushes and 
small trees as well as mangroves (Taylor. 1922d; Lev- 
iton. 1964b; Alcala, 1986). The Mt. Madja-as speci- 
men was found in a banana plantation at 800 m near 
primary forest. We find it unlikely that all SE Asian 
populations currently referred to T. wagleri will prove 
to be a single species. This is the first account of a 
specimen referable to this species from Panay. 

Localities and specimens: (Site 1 1 ) CMNH 5076. 



Discussion 

The primary goal of this report has been to provide a 
comprehensive and synthetic review of the amphibian 
and reptiles species known from Panay Island and is 
surrounding land-bridge islets. As noted, over the past 
century, biologists have expected that the documented 
presence of a species on the neighboring island of 
Negros implied its undocumented presence on Panay 
as well (see Inger, 1954; Brown and Alcala. 1970. 
1978, 1980. 1986; Brown and Rabor, 1967: Alcala. 
1986). However, Panay (and its smaller satellite islets) 
supports low levels of endemicity, primarily as exem- 
plified by populations at higher montane elevations. 
Panay has several species of amphibians and reptiles 
that have not yet been reported on Negros or Cebu, is 
known to support species that so far have not been 
documented on Panay. The presence on Panay of 
numerous new and undescribed species suggests that 
calculations of Panay 's herpetological endemicity will 
continue to rise with continued survey work. Addi- 
tionally, basic taxonomic studies of species com- 
plexes with representatives on Panay will no doubt 
further contribute to the total number of Panay 
endemics. 

There are only a few reports of amphibians and 
reptiles of high elevation forests of Panay (Sison et 
al., 1995; W Brown et al.. 1997a; Ferner et al., 1997; 
R. Brown et al. 1999, this issue). Essentially, on each 
occasion that herpetologists have surveyed higher ele- 
vation forests of Panay, they have discovered new spe- 
cies. Other surveys conducted in forested regions of 
Panay include survey efforts of A. Diesmos, R. Crom- 
bie. and M.GauIke (in press). Further high elevation 
surveys in well forested regions of Panay are greatly 
needed to gain an understanding and appreciation of 
these presumably relictal faunal elements. 

There are numerous records that are included in 
this report that were not at all unexpected. These 
include widespread SE Asian and Philippine species 
that biologists have expected or assumed were present 
on Panay. Others have even been listed as known from 
Panay, but without specific reference to reliable local- 
ity data or museum specimens (see Alcala. 1986). 
These include the frogs Bufo marinus, Kaloula picta, 
Occidozyga laevis, Platymantis corrugatus, Platy- 
mantis dorsalis, Rami vittigera, the skink Mabuya 
multifasciata, and the snakes Ramphotyphlops brami- 
nus, Acrochordus granulatus, Psammodynastes put- 
verulenius. Gonyosoma oxycephala, and 
Tropidolaemus cf wagleri. 

Another group of species includes forms known 
from other well-surveyed islands in the Visayas 



Vol. 9, p. 56 



Asiatic Herpetological Research 



2001 





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2001 



Asiatic Herpetological Research 



Vol. 9. p. 57 






Vol. 9, p. 58 



Asiatic Herpetological Research 



2001 




Fig. 1 1 . Flooded mangrove forest on 
northeast coast of Negros Island (Photo: 
RMB). 





Fig. 12. South coast of Gigante North 
Island (Site 35; Photo: RMB). 




Fig. 13. Limstone cave on Gigante South 
Island; habitat of Gekko gigante (Site 37; 
Photo: RMB). 



Fig. 14. Jagged karst limestone habitat of 
Platymantis insulatus on Gigante South 
Island (Site 36; Photo: RMB). 



2001 



Asiatic Herpetological Research 



Vol. 9. p. 59 




-ig. 15. Bufo marinus (Photo: RMB). 



Fig. 16. Kaloula corijuncta negrosensis (Photo: 
RMB). 







ig. 17. Kaloula picta (Photo: RMB). 



Fig. 18. Limnonectes cf leytensis (Photo: RMB). 




: ig. 19. Limnonectes visayanus (Photo: RMB). Fig. 20. Occidozyga laevis (Photo: RMB). 




: ig. 21 . Platymantis corrugatus (Photo: RMB). Fig. 22. Platymantis insulatus (Photo: RMB). 



Vol. 9, p. 60 



Asiatic Herpetological Research 



2001 




Fig. 23. Rana erythraea (Photo: RMB) 



Fig. 24. Rana vittigera (Photo: RMB). 




Fig. 25. Polypedates leucomystax (Photo: 
RMB). 



Fig. 26. Cuora amboinensis (Photo: RMB). 

m 




Fig. 27. Draco spilopterus (Photo: J. McGuire). Fig. 28. Draco spilopterus with extended 

patagium (Photo: J. McGuire). 




Fig. 29. Mature female Hydrosaurus pustulatus Fig. 30. Immature female Hydrosaurus 
(Site 23; Photo C. Banks). pustulatus (Site 1 1 ; Photo; J. McGuire). 



2001 



Asiatic Herpetological Research 



Vol. 9. p. 61 





<~ ~*-t ~X.4 







Fig. 31 . Mature male Gonocephalus sp (Photo: 
RMB). 



Fig. 32. Cyrtodactylus annulatus (Photo: RMB). 




Fig. 33. Cyrfodacfy/us philippinicus (Photo: 
RMB). 



Fig. 34. Gehyra mutilata (Photo: RMB). 











^r 


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A. ^^k 






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Fig. 35. 



gigante (Site 37: Photo RMB) 




Fig. 36. Gekko gigante eggs in cave crevice 
(Site 37; Photo: RMB). 



~» - «* 






- > V~M^^; 




Fig. 37. Gekko mindorensis (Photo: RMB) 




Fig. 38. Brachymeles boulengehtaylon (Photo: 
RMB). 



Vol. 9, p. 62 



Asiatic Herpetological Research 



2001 




-■• " -«p 



Fig. 39. Brachymeles boulengeri taylori (Photo: Fig. 40. Brachymeles tridactylus (Photo: RMB). 
RMB). 




Fig. 41 . Mabuya multicarinata borealis (Photo: Fig. 42. Sphenomorphus jagori grandis (Photo: 
J. McGuire). ' RMB). 




Fig. 43. Tropidophorus grayi (Photo: RMB). 



Fig. 44. Varanus salvator nuchalis, dorsal view 
(Photo: J. McGuire). 




Fig. 45. Varanus salvator nuchalis, ventral view Fig. 46. Python reticulatus (Photo: RMB). 
(Photo: J. McGuire). 



2001 



Asiatic Herpetological Research 



Vol. 9. p. 63 




Fig. 47. Ahaetulla prasina preocularis (Photo 
RMB 

I ■■ 



Fig. 48 Dendrelaphis pictus pictus (Photo: 
RMB). 




Fig. 49. Lycodon aulicus capucinus (Photo: 
RMB). 




Fig. 50. Pseudorabdion mcnamarae (Photo: 
RMB). 




Fig. 51 . Typical male Tropidolaemus cf wagleri 
(Photo: RMB). 



Fig. 52. Typical female Tropidolaemus cf 
wagleri (Photo: RMB). 






Vol. 9, p. 64 



Asiatic Herpetological Research 



2001 



(Negros. Cebu) that we expected to find on Panay as 
well. Their presence on Panay was almost a certainty 
based on known biogeographic relationships of these 
islands. These species include the frogs Kaloula c. 
negrosensis, Limnonectes cf leytensis. the skinks 
Lipinia pulchella taylori, Brachymeles boulengeri 
taylori, Mabuya multicarinata borealis, Tropidopho- 
rus grayi. Emoia atrocostata, the geckos Cyrtodacty- 
lus annulatus, Cyrtodactylm philippinicus, the sail-tin 
agamid Hydro saurus pustulatus, and the snakes Boiga 
angulata. Boiga cf cynodon, Boiga cf dendophilia, 
Cyclocorus lineatus alcalai, Pseudorabdion mcnama- 
rae, Oligodon modestum, and Trimereserus flavomac- 
ulatus (Gaulke, in press). 

Several other records are major range extensions 
and real surprises. The skink Dasia semisincta is oth- 
erwise known only from Borneo and Mindanao Island 
and its presence on Panay is based on a well 
vouchered specimen (USNM 78840) that can not be 
discounted. The presence of a new species in the 
genus Hologerrhum was surprising in that this genus 
previously was considered a monotypic Luzon Aggre- 
gate Island Complex endemic (Leviton, 1963c; Brown 
et al., this issue; Gaulke. in press). A forest species of 
Kaloula related to K. kalingensis and K. kokacii is the 
first record of this species group outside the bound- 
aries of the Luzon Aggregate Island Complex (Inger, 
1954; Brown and Alcala. 1970; Alcala and Brown, 
1998; Brown and Diesmos, unpublished data; Gaulke. 
in press). Some records (based on few specimens or 
dubious locality data) may be in error: these include 
Sphenomorphus coxi divergens, Sphenomorphus cum- 
ingi, and S. fasciatus (of Sison et al., 1995); other- 
wise, if accurate, these records represent major range 
extensions beyond the confines of the Mindanao. 
Mindoro. and Luzon aggregate island platforms. 

As mentioned, a number of new species recently 
have been described as Panay endemics; the presence 
of endemics in high elevation habitats of Panay was 
not unexpected. These species Hologerrhum dermali 
(Brown et al.. this issue; see also Gaulke, in press). 
Parvoscinus sisoni (Ferner et al., 1997). Platymantis 
panaxensis (Brown et al., 1997a), Pseudorabdion tal- 
onuran (Brown et al., 1999). Kaloula sp., Platymantis 
sp. 1, Platymantis sp. 2. and Platymantis sp. 3 (this 
report, see also Gaulke, in press). 

Finally, there are numerous "subspecies", species, 
or members of widespread Visayan. Philippine, or SE 
Asian species complex members on Panay that are of 
uncertain taxonomic status and in need of basic taxo- 
nomic review (see individual species accounts). Many 
species currently listed from Panay are. we expect, 
distinct lineages that eventually will be recognized as 



Panay or Visayan endemics. These species are too 
numerous to list and extend from common, low eleva- 
tion forms to rare high elevation forest obligates. 
There is much basic taxonomic work still to be con- 
ducted on Panay. in the Visayas. and in the rest of the 
Philippines and we expect our estimates of Panay 's 
endemicity to generally rise with further systematic 
studies of the herpetofauna of the Philippines. 

Faunal similarity calculations (Fig. 53) indicated 
that, as expected, the Visayan islands of Negros and 
Cebu (situated on the same Pleistocene aggregate 
island platform as Panay) were among the islands fau- 
nistically most similar to Panay (Masbate was not 
considered due to the absence of substantial records 
from this island). We interpret this as evidence for 
mid- to late-Pleistocene land bridge connections 
between these islands (see Heaney, 1985. 1986). 
Within the Visayan Aggregate Island Complex, esti- 
mates of amphibian faunal similarities exceeded those 
of reptiles but when Panay is compared to islands out- 
side the Visayan Aggregate Island Complex, reptile 
faunal similarities exceed those of amphibians. As 
expected, amphibian faunal similarity between Luzon 
and Panay is much lower than estimates calculated for 
Cebu or Negros and Panay. However, surprisingly, 
Luzon and Panay had a higher reptile coefficient of 
similarity than did Cebu and Panay. A portion of this 
unusual finding may be the result of sampling error 
reflecting the degree to which survey data for Negros. 
Panay. and Luzon are available. Calculations of simi- 
larities between Negros and Luzon (not shown) are 
higher in reptiles (0.58) than they are for amphibians 
(0.45). 

In comparisons with islands on other (non- 
Visayan) major Philippine aggregate island platforms, 
reptilian species similarity was consistently higher 
than that of amphibians. This may in part be due to 
antiquated taxonomy. Recently, disproportionate 
amounts of taxonomic work has been conducted on 
amphibian groups, resulting in the recognition of 
more amphibian endemics, with fewer "shared" spe- 
cies among islands. In contrast, many reptiles "spe- 
cies" are shared between Panay and Luzon. Some of 
these may represent species complexes in need of tax- 
onomic resolution. Alternatively, these calculations 
may reflect the relatively greater dispersal abilities 
presumed for reptiles when crossing ocean barriers. It 
is tempting to consider that more reptile species may 
be shared between these islands because reptiles may 
be more tolerant of exposure to heat and salt water, 
and may have a higher probability of surviving dis- 
persal events (e.g.. via rafting) than would be 
expected for amphibians. In general, these results sup- 



2(K)1 



Asiatic Herpetological Research 



Vol. 9. p. 65 




Figure 53. Faunal similarity coeffi cients, calculated bet 
ween Panay and other major islands in the Philippines. 
See text for formula and discussion. 

port the suggestion that the herpetofaunal communi- 
ties of islands within the Visayan Aggregate Island 
Complex are very similar but also that they have their 
own degree of endemism and are far from being iden- 
tical. 

Species that we might expect to soon be discov- 
ered on Panay include populations that are otherwise 
known from Negros and Cebu islands. However, we 
note that the high elevation endemics of Negros (e.g., 
Platymantis hazelae, Pseudorabdion montanum) are 
not expected to be discovered on Panay; in their place 
we expect Panay to support it's own high elevation 
populations of closely-related montane endemics 
(e.g., Platymantis panayensis, Pseudorabdion talonu- 
ran). These include the frogs Rhacophorus pardalis 
(Fig. 54). Rhacophorus appendiculatus (widespread 
on Mindanao and Luzon aggregate island complexes: 
both known from Negros), Platymantis spelaeus 
(known from Negros; Fig. 55). geckos such as Lepi- 
dodactylus lugubris (widespread in the Philippines), 
Lepidodactylus herrei (currently comprised of two 
subspecies: L. h. herrei on Negros and L. h. medianus 
on Cebu), Lepidodactylus christiani (known from 
Negros and Cebu). Pseudogekko brevipes (known 
from Negros and Cebu: P. compressicorpus has been 
collected on Masbate). a Luperosaurus species (L. 
cumingi has been collected on Negros), and skinks 
like Lipinia quadrivittata quadrivittata (from Negros 
and Cebu). Lipinia auhculata auriculata (from 
Negros and Masbate), and Lipinia rabori (from 
Negros). Snake species we expect will be found on 




Fig. 54. Rhacophorus pardalis, present on 
Nergos but not yet recorded on Panay (Photo: 
RMB). 




1 



Fig. 55. Platymantis spelaeus, known from 
caves in southern Negros: this species has not 
yet been recorded on Panay (Photo: RMB). 




Fig. 56. Oxyrhabdion leponnum visayanum. 
well-known and common throughout Nergos 
Island, but not yet recorded on Panay (Photo: 
RMB). 

Panay with continued survey efforts include forms 
possibly related to T. canlaonensis, T. hedraeus 
(Negros forms). Oxyrhabdion leponnum visayanum 
(from Negros and Cebu; Fig. 56). and Ophiophagus 
Hannah (recorded from numerous islands in the Phil- 
ippines). The Philippine endemic crocodile. Crocody- 
lus mindorensis. may have recently been rediscovered 



Vol. 9, p. 66 



Asiatic Herpetological Research 



2001 



on Negros (E. Alcala, personal communication) and 
may be present on Panay if suitable habitat can be 
located. As noted, the Gigante Island endemics Plan- 
mantis insulatus and Gekko gigante might be 
expected to occur on karst limestone outcrops on 
Panay's northeast coast. 

The future of exploration on Panay guarantees 
continuation of the kind of discovery reported here. 
There is an immediate need for continued basic sur- 
vey efforts in the montane portions of Antique. Aklan. 
and Iloilo provinces, all of which contribute to the 
western coastal mountain range that supports so much 
of Panay's herpetological endemicity. Additionally, 
low elevation portions of Panay (principally Capiz 
and Iloilo provinces) are also herpetologically 
unknown. If areas of overlooked primary forest or 
well-regenerated secondary forest can be located, we 
have high expectations that these will support novel 
herpetological communities and generate continued 
discoveries of new taxa. Areas of particular interest 
include karst limestone outcrops along the northern 
and eastern coasts, mangrove fragments, cave habi- 
tats, and isolated outcrops of moderate elevation in 
eastern Panay (Fig. 2). As noted, the best place to 
search for the Gigante endemics Platymantis insula- 
tus and Gekko gigante is limestone outcrops support- 
ing caves along Panay's northeastern coast. Other rare 
Negros species (i.e.. Luperosaurus cumingi, Lepido- 
dactylus herrei, Platymantis spelaeus) may eventually 
be revealed on Panay as well, once adequate surveys 
in preferred microhabitats (forest canopies and karst 
limestone caves; C. N. Dolino, personal communica- 
tion) become available. 

Like most islands in the Visayan Aggregate Island 
Complex. Panay should be regarded a priority for 
future conservation initiatives and programs aimed at 
sustainable resource management. Panay is a unique 
island (not at all identical to Negros) that deserves its 
own conservation efforts. 

The few remaining forests of Panay continue to be 
felled at an alarming rate, suggesting that its endemic 
flora and fauna may disappear before even being 
recorded by biologists. Low elevation forests and 
mangroves are all but gone, and even the most dis- 
turbed and negatively impacted sites warrant immedi- 
ate study of the kind that currently is underway on 
Cebu and Negros (A. Alcala. and E. Alcala, C. N. 
Dolino. J. C. Gonzales, and M. Pedregosa. personal 
communication). It is our hope that Panay will be rec- 
ognized as a model island ecosystem, ripe for collabo- 
rative efforts of conservation biologists, taxonomists. 
biogeographers, community organizers, and politi- 
cians. Conservation efforts targeted at the community 



level represent the best opportunity for foreign, gov- 
ernment, and non-government organizations in their 
effort to halt the destructive practices of non-sustain- 
able timber and mineral extraction industries that cur- 
rently operate unchecked in the central Visayan 
islands of the Philippines. 

Acknowledgements 

We thank the management and staff of the Protected 
Areas and Wildlife Bureau of the Philippines Depart- 
ment of the Environment and Natural Resources 
(DENR) for facilitating collecting and export permits 
necessary for portions of this study. For logistical 
assistance, we thank P. Gonzales (PNM) and M. 
Ebreo (DENR, Iloilo City), the officers and staff of 
the Regional DENR offices in Iloilo, and the provin- 
cial DENR authorities of the Municipalities of Valder- 
rama. Culasi. and San Jose. The PNM/CMNH 
Philippine Biodiversity Inventory (PBI) was funded in 
part by a grant from the John D. And Catherine T. 
MacArthur Foundation, with additional support from 
CMNH benefactors. Support for JWF's Panay field 
work (1989 and 1992) came from CMNH and the 
Faculty Development Fund and Department of Biol- 
ogy. Thomas More College. Support for RMB's par- 
ticipation in field work on Panay (1992. 1993, 1996) 
came from the Roschman Student Enrichment Fund, 
the Alumni and Friends of Miami University Under- 
graduate Research Grant. Zoology and Botany 
Departments, and the College of Arts and Sciences of 
Miami University, and CMNH. We thank the follow- 
ing individuals (and their respective institutions) for 
assistance, provision of working space, and help 
assembling museum records: J. Vindum (CAS). A. 
Resetar (FMNH). R. Crombie (USNM). and J 
Rosado (MCZ). Financial support for RMB's travel to 
CAS was provided by the Stearns Grant of the Cali- 
fornia Academy of Sciences. The dedicated assis- 
tance, untiring enthusiasm, and cheerful company of 
K. Auffenberg. J. Barcelona, J. Bulalacao. D. Burt, D. 
Busemeyer. J. Cabalquinto, E. Canada, P. Comintan. 
A. Diesmos, J. Demboski, M. Ebreo, R. Fernandez. 
J.Lasugas. M. Manuel. J. McGuire. H. Miranda, E. 
Mockford. L. Moores, K. Reis.L and D. Ruedas, J. 
Ruthven. E. Sagcal. V Samarita. R. Wacdisen. and V. 
Yngente is greatly appreciated. We owe particular 
thanks to A. Alcala for his effort to organize and logis- 
tically support a recent excursion to Gigante Island. 
We thank J. Barcelona, L. Bockstanz, D. Cannatella, 
R. Crombie. A. Diesmos, M. Gaulke, J. McGuire, and 
J. Slowinski for comments on portions of earlier 
drafts of this paper, and R. Crombie and M. Gaulke 
for sharing their ideas, time, and unpublished data. 



2001 



Asiatic Herpetological Research 



Vol. 9. p. 67 



This paper is contribution no. 27 to the results of the 
PNM/CMNH PBI. 

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106:34-45. 



2001 



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Vol. 9. pp. 71-76 



The Discovery of Mauremys iversonh\ ike Turtles at a Turtle Farm in Hainan 
Province, China: The Counterfeit Golden Coin 

James Ford Parham 1 and Haitao Shi 2 

Department of Integrative Biology, University of California, Berkeley, California 94720-3140, USA, email: 

parham@socrates.berkeley.edu. Department of Biology, Hainan Normal University, Haikou City, Hainan 

Province 571 158, China 

Abstract.- During a visit to a turtle farm in Hainan Province, we discovered turtles that closely resemble 
Mauremys iversoni. The Mauremys iversoni-hke turtles at the turtle farm were intentionally produced hybrids of 
two commonly reared species, Mauremys mutica and Cuora trifasciata. According to the turtle farmer, the 
intentional production of hybrids between these two turtles is a common practice. The hybrids fetch extremely 
high prices (-1000 USD/kg) since they are sold as Cuora trifasciata. the "golden coin" turtle, to unsuspecting 
buyers. We suggest, but cannot prove, that all Mauremys iversoni-Mke turtles might be turtle farm hybrids. This 
hypothesis could explain all of the available evidence and could alleviate a lot of the confusion surrounding this 
species unusual geographical distribution, sudden appearance in the pet trade, as well as its variable morphology. 

Key words.- Turtles, Bataguridae, Geoemydidae. Mauremys. Cuora trifasciata. Mauremys iversoni. China. 
Hainan, hybrids, turtle farm 



Introduction 

Mauremys iversoni Pritchard and McCord 1991 is one 
of ten new Chinese chelonians described from speci- 
mens purchased through the pet trade since 1987. 
From the beginning, our knowledge of this species' 
geographical distribution has been fraught with confu- 
sion. The type specimen of Mauremxs iversoni (UF 
71866: institutional abbreviations follow Leviton et 
aL 1985) and 19 other individuals are reported to 
have come from Fujian province (Fig. la), but eight 
additional specimens from a locality in Guizhou (over 
1000 km away) were also reported (Fig. lb). Prit- 
chard and McCord ( 1991 ) propose two hypotheses to 
explain this disjunct distribution: 1) The distribution 
is (or was) continuous between these localities; 2) 
One of the localities (perhaps the type locality) is the 
result of turtles being relocated by traveling Bud- 
dhists. 

An additional wrinkle is provided by Iverson and 
McCord (1994) who suggest that the type series of 
Mauremys "guangxiensis" (-'Clemmys guangxien- 
sis" Qin 1992) from Guangxi (Fig. lc) is actually a 
composite of Mauremys mutica and Mauremxs iver- 
sow- like specimens. However, some differences 
between the M. "guangxiensis" iversoni-Mke speci- 
mens, the Guizhou M. iversoni specimens, and the 
Fujian M. iversoni specimens have been noted (Prit- 
chard and McCord. 1991; Iverson and McCord, 
1994). The result is the occurrence of different-look- 
ing M. iversoni-\ikc turtles from three disparate 




Figure 1. The reported localities of Mauremys iversoni 
like turtles in China: A) Type locality, Nanping, Fujian 
Province (Pritchard and McCord, 1991): B) Kweiying, 
Guizhou Province (Pritchard and McCord, 1991); C) 
Nanning, Guangxi Province (Qin, 1992); D)Tunchang, 
Hainan Province (This study). 

regions in China between the years of 1986 and 1991 
(Fig. 1 ). Based on observations from a breeding facil- 
ity located on Hainan Island (Fig. Id), we propose a 
third hypothesis that might explain all of the available 
data and possibly alleviate much of the confusion: All 
Mauremxs iversoni-\'\ke turtles are the result of the 



Vol. 9, p. 72 



Asiatic Herpetological Research 



2001 




Figure 2. An intentionally produced hybrid trom a Chi- 
nese turtle farm (MVZ 230475) that closely resembles 
Mauremys iversoni. Photo by JFR 

hybridization of Mauremys mutica and Cuora trifasci- 
ata in captivity. 

Chinese turtle farms 

The role of turtle farms in the Asian turtle trade is 
often overlooked. The primary reason for this is the 
secrecy of the turtle farmers themselves. Despite our 
ignorance, the practice of farming turtles in China is a 
widespread, lucrative endeavor. As far back as 1991, 
Zhou and Zhou report that Cuora trifasciata is being 



bred "everywhere". In the past ten years, the number 
of turtle farms is rumored to have increased dramati- 
cally. Although it is not possible to provide exact 
numbers, there are estimated to be at least ten breed- 
ing facilities on Hainan alone. In an attempt to assess 
the impact of this burgeoning business to the Asian 
turtle trade, one of us (HS) has gained access to a 
large turtle farm in Tunchang. Hainan province (Fig. 
Id) from 1996 to the present. This paper represents 
the second report of this effort (see also Shi and Par- 
ham, 2001). 

According to the owner of the turtle farm, the Tun- 
chang breeding facility was first established in 1983. 
The foundation of its breeding stock came from doz- 
ens of wild collected Cuora trifasciata. Mauremys 
mutica, and Ocadia sinensis from Hainan as well as 
additional specimens from a farm in Guangdong that 
was established in 1978. It currently houses an esti- 
mated 15.000+ geoemydids. Early on. the number of 
breeding ponds and turtles was limited and the estab- 
lishment of breeding stocks proceeded without clear 
aim. Almost all the local species of turtles were col- 
lected from the field or bought from villagers and 
often kept in the same pond or enclosure. 

In November of 1999, the authors visited a smaller 
facility that included several indoor breeding ponds. 




Figure 3. A comparison of Mauremys iversonip\as\ta from the type description with turtle farm hybrids and putative 
parental species: A) Mauremys mutica Uom a turtle farm in Tunchang, Hainan Province (MVZ 230477); B-D) Mau- 
remys tVerson/Uom the type description. They are either from Fujian or Guizhou Province; E-F) Intentionally pro- 
duced hybrids from a turtle farm in Tunchang, Hainan Province; G) Mauremys iverson/'Uom the type description. It 
is either from Fujian or Guizhou Province; H) An intentionally produced hybrid from a turtle farm in Tunchang, 
Hainan Province (MVZ 130475); I) Cuora trifasciata 'from Tai Pin, central Hainan Province (MVZ 23932); J) Cuora 
trifasciata Atom the pet trade (MVZ 230636). Photos by JFP [a,e-f,h-j] and Pritchard and McCord (1991 )[b-d,g]. 



2001 



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Vol. 9, p. 73 



Although numerous species were observed, Maure- 
mys mutica and Cuora trifasciata were the most abun- 
dant. Many of the C. trifasciata were kept separately. 
but one pond included C. trifasciata. M. mutica. and 
even a Chelydra serpentina. When asked if any of the 
turtles ever hybridized, the workers of the farm pro- 
duced several animals that closely resembled Maure- 
mys iversoni. One of these hybrids, a subadult male, 
was procured as a voucher (MVZ 230475; Fig. 2. 3h). 
JFP was given permission to photograph two other 
hybrids (Fig. 3e,f). 

The turtle farm hybrids 

As with the three previously reported incidences of M. 
iversoni (Fujian, Guizhou. and Guangxi). there are 
some differences in the Hainan farm specimens. 
Unfortunately, it is difficult to understand these differ- 
ences because of inconsistencies in the reporting of 
previous measurements, small sample sizes, and the 
fact the most important specimens remain in private 
hands. For example, Pritchard and McCord (1991) 
present bivariate plots of measurements for adult 
Mauremys iversoni. but inexplicably include only five 
of the adult males. It is not stated from which locality 
(Fujian or Guizhou) these specimens are from or why 
the measurements of all 10 of the adult males from 
Fujian (the type locality) are not included. Table 1 of 
Pritchard and McCord includes 10 shell measure- 
ments for all the specimens, but excludes the two 
measurements (the interpectoral length [IPLj and the 
interanal length [IAN]) that would have allowed the 
type series to be included in the bivariate plot. Iver- 
son and McCord (1994) perform a more detailed 
study of variation within East Asian Mauremys. but do 
not present the raw data and only include one speci- 
men of Guizhou Mauremys iversoni (no voucher). 

The relevant shell measurements of MVZ 230475 
are as follows: IAN= 1.4 cm, IPL= 1.1 cm. CL= 1 1.4 
cm. IAN/CL= .1228, IPL/CL=.096. MVZ 230475 is 
a juvenile and smaller than three specimens excluded 
from the adult category by Pritchard and McCord 
(1991). Bearing this consideration, a comparison 
shows that the morphology of MVZ 230475 is unlike 
the type series of Mauremys iversoni. and apparently 
unlike the other Mauremys. It is uncertain what the 
variation between and among the Mauremys iversoni 
series (from Guizhou and Fujian) is fully represented 
by the polygon in Pritchard and McCord (1991). 
Since the relevant scale ratios from their entire series, 
including the subadult specimens, are not reported, 
we are forced to rely on the close phenetic similarity 
between the turtle farm hybrids and M. iversoni for 
our conclusions. It is important to emphasize that if 




Figure 4. Cuora trifasciata Uom a Chinese turtle farm 
showing light head coloration. Photo by HS. 

all M. iversoni are turtle farm hybrids, then we should 
expect them to have an extremely variable morphol- 
ogy, especially if they are the result of separate hybri- 
dogenic events. This pattern is borne out by the 
previously reported M. iversoni-\\kc turtles (Pritchard 
and McCord, 1991; Qin. 1992; Iverson and McCord, 
1994). 

In terms of coloration, MVZ 230475 and the 
hybrids photographed at the Tunchang turtle farm 
closely resemble Mauremys iversoni. The head color- 
ation of MVZ 230475 is lighter than that of the speci- 
men figured by Pritchard and McCord (1991 ). but a 
variation in head coloration in Mauremys iversoni was 
noted by Fritz and Obst (1999). MVZ 230475 is 
almost identical to the lighter individual figured in 
that work. The variable head coloration in M. iversoni 
can be linked to the variation in C. trifasciata. For, 
while most C. trifasciata have dark postorbital mark- 
ings. C. trifasciata with light head coloration are 
known (Fig. 4), especially in turtle farms. Pritchard 
and McCord ( 1991 ) state that M. iversoni can be dis- 
tinguished from M. mutica by a horseshoe-shaped 
coalescence of blotches from the pectorals to the anals 
(with the open end anterior). Although they state that 
this pattern is not found in Mauremys mutica. Zhou 
and Zhou ( 1991, p. 38) illustrate a M. mutica (locality 
not stated) with this this plastral pattern. Later. Iver- 
son and McCord (1994) illustrate a M. mutica from 
Taiwan (FMNH 127181 ) that also has a clear horse- 
shoe-shaped coalescence of blotches. Even without 
this character, M. iversoni can be distinguished from 
M. mutica by its more rounded anterior lobe that lacks 
a prominent gular projection. Furthermore, many M. 
iversoni specimens, including some of the type series 



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Asiatic Herpetological Research 



2001 



of M. iversoni and the Hainan farm hybrids (Fig. 3b- f) 
have transverse trending blotches on the pectorals, a 
feature not known to occur in Mauremys, but common 
in juveniles of C. trifasciata. 

The variation among the figured specimens in the 
type description is extreme (Fig. 3b-d,g). It is not 
stated whether these specimens are from Fujian or 
Guizhou. Comparisons of the plastral figures from 
Pritchard and McCord (1991) with the specimens 
from the Tunchang farm show that the latter speci- 
mens lie within the range of variation of M. iversoni. 
The most notable differences are that one (Fig. 3e) has 
more irregularly shaped plastral blotches and MVZ 
230475 (Fig. 3h) has only a narrow, light, central fig- 
ure on the plastron. Perhaps the most important fea- 
ture to be noted is that no two specimens look alike. 
In the type description, Pritchard and McCord ( 1991 ) 
describe the plastral pigmentation as "very variable in 
intensity", but it is unclear whether the figured speci- 
mens represent the most typical patterns of Mauremys 
iversoni or the entire range of variation. 

Luckily, there are three characters of M. iversoni 
that clearly set it apart from other East Asian Maure- 
mys: 1) its olive or yellow head, 2) rounded end of 
anterior lobe of the plastron and 3) red coloration on 
the underside of the limbs. Until the description of 
Mauremys iversoni, the red coloration of the limbs 
was a diagnostic characteristic of C. trifasciata. The 
light head coloration and rounded lobe are also remi- 
niscent of C. trifasciata. Furthermore, in Mauremys 
iversoni the sulcus between the humeral and abdomi- 
nal scutes lies closer to the hyo/hypoplastral suture 
than it does in other Mauremys. An alignment of this 
sulcus with the junction of these bones is a character 
shared by C. trifasciata and other taxa with a kinetic 
plastron. In other words, the morphology of Maure- 
mys iversoni is intermediate between C. trifasciata 
and M. mutica, and therefore consistent with a hybrid 
origin. A detailed comparison of the mitochondrial 
DNA and allozymes of this specimen and other pet 
trade "species" is currently under way (Parham et al., 
2000; in prep.). 

Discussion 

During our joint visit to the Tunchang turtle farm, the 
workers stated that production of Mauremys iversoni- 
like turtles was the result of infrequent, accidental 
hybridization events. Since that time, the owner of 
the farm has confided that the intentional production 
of C. trifasciata X M. mutica hybrids is a common 
practice undertaken by several turtle farmers. 
Because practitioners of Chinese traditional medicine 
claim that C. trifasciata has many medicinal proper- 



ties, and recently it has also been suggested that it has 
the ability to cure cancer, it is highly valued. As a 
result, it is often called the "golden coin" turtle. The 
price of turtles is determined by their weight. In the 
year 2000. 1 kg of M. mutica sold for approximately 
100-120 yuan (-15 USD) whereas 1 kg of C. trifasci- 
ata sold for 6,000-8,000 yuan (-1000 USD). Accord- 
ing to the turtle farmer, it is possible to sell the 
hybrids of C. trifasciata and M. mutica as pure C. tri- 
fasciata since, to the untrained eye, these species 
closely resemble one another, especially in the yellow 
coloration of the head. The confusion of these two 
forms should not be surprising since Timmins and 
Khounboline (1999) report that even people familiar 
with C. trifasciata have mistakenly identified juvenile 
M. mutica as this species. Therefore, it is possible to 
produce and sell large numbers of counterfeit "golden 
coins" to unsuspecting buyers without having to 
obtain large numbers of adult Cuora trifasciata. 

The high degree of phenetic similarity between 
the C. trifasciata X M. mutica hybrids from Hainan 
and the M. iversoni-tokc turtles reported from Fujian, 
Guizhou and Guangxi strongly suggests that all of 
these animals may be the product of accidental or 
intentional hybridization in turtle farms. This hypoth- 
esis explains the unusual morphology, its sudden 
appearance in the pet trade, absence in historical col- 
lections, and the confusion surrounding the distribu- 
tion of this taxon. The timing of the discovery of 
these turtles shortly follows the increased demand for 
Cuora trifasciata as a reputed cure for cancer (van 
Dijk et al., 2000). This increased demand may have 
initiated the establishment of numerous turtle breed- 
ing facilities that led to the production of Mauremys 
iversoni-Yikc turtles. 

We stress that the practice of housing and breeding 
several species of chelonians in a single enclosure or 
pond is probably not restricted to the Tunchang farm 
alone. If the establishment of other facilities mirrors 
the one on Hainan, the sudden appearance of new and 
unusual turtles in the pet trade would be expected. 
The ability of turtles to hybridize is well documented 
(Fritz and Baur, 1994; Fritz, 1995). Certain species, 
such as Mauremys pritchardi McCord 1997. have 
already been implicated as possible hybrids (Artner et 
al.. 1998). Others, such as Sacalia pseudocellata 
Iverson and McCord 1992. Ocadia glyphistoma 
McCord and Iverson 1994. and Ocadia phillipeni 
McCord and Iverson 1992 are known from a paltry 
number of specimens which might be indicative of an 
accidental production in a turtle farm (van Dijk, 2000; 
Lau and Shi. 2000). It is probably not a coincidence 
that many of the newly described species (O. phil- 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 75 



lipeni, S. pseudocellata, and M. iversoni) can be dis- 
tinguished from their congeners by characters found 
in the commonly reared and highly valued C. thfasci- 
ata. Studies of the DNA of the holotypes of these 
species, compared with multiple, field-collected, 
vouchered specimens of established species from 
known localities, could easily determine the validity 
of these taxa. 

Conclusions 

It is uncertain whether all of the reported collection 
data from the pet trade, such as the distributional data 
for Mauremys iversoni are actual or fabricated. The 
rest of the available evidence strongly suggests, but 
does not prove, that all M. iversoni-Wke turtles could 
have a turtle farm origin. Parham and Li (1999) 
openly question the Yunnan locality of Cuora pani 
that was provided by the same pet dealer credited with 
collecting the original Mauremys iversoni series. The 
high prices that unusual turtles with locality data can 
fetch from turtle fanciers could serve as incentive to 
fabricate collection data. Furthermore, there is also 
incentive to hide the true locality as a trade secret, 
especially if the locality is a Chinese turtle farm. Of 
course, given the current scarcity of turtles in Asia it is 
difficult to prove that a locality is in error or whether 
the turtles have been extirpated. Verified distribution 
data for rare chelonians (e.g., Iverson, 1992) are criti- 
cal to understanding their current and historical status. 
False localities unnecessarily complicate our meager 
understanding of turtle distributions. In light of the 
confusion surrounding pet trade data, we recommend 
that workers should treat them with utmost caution 
until they are supported by evidence that is more reli- 
able. 

Given the seemingly uncontrollable Asian Turtle 
Crisis, determining which species are valid evolution- 
ary lineages and which are turtle farm hybrids is 
important. . Otherwise, crucial resources could be 
wasted on invalid taxa. For example, the Red data 
Book of Endangered Animals (Wang, 1998) suggests 
that Fujian and Guizhou provinces protect M. iversoni 
and that surveys and ecological studies should be per- 
formed in order to determine the proper conservation 
strategies. We concur with van Dijk's (2000) asser- 
tion that researchers should make the investigations 
on the validity of the pet trade species a priority so 
that the limited resources for conservation can be 
directed towards established taxa that are undergoing 
unimagined, precipitous declines. 



Acknowledgments 

The authors would like to thank John Iverson for 
prompting a comparison between the turtle farm 
hybrids and the type description of Mauremys iver- 
soni. We would also like to thank Dr. Jaeger, the edi- 
tor of Herpetologica, for granting us permission to 
reproduce part of Figure 1 from Pritchard and 
McCord (1991). Ted Papenfuss and Jim Buskirk pro- 
vided helpful comments. JFP is also grateful to Carl 
Gillies, Kevin Padian, and Tonya Van Leuvan-Smith. 
This project was funded by the National Science 
Foundation (JFP), the University of California 
Museum of Paleontology (JFP), The National Natural 
Science Foundation of China (HS), The Provincial 
Natural Science Foundation of Hainan (HS), and the 
Hainan Provincial Ecological Key Discipline Fund 
(HS). 

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A Review of the Distribution of Diploid, Triploid andTetraploid Green Toads 
{Bufo viridis Complex) in Asia Including New Data from Iran and Pakistan 1 

Matthias Stock 1 , Daniel Frynta 2 , Wolf-Rudiger Grosse 1 , Claus Steinlein 3 and 

Michael Schmid 3 

Martin-Luther-Universitdt Halle-Wittenberg, Institutfiir Zoologie, Domplatz 4, D - 06099 Halle/ Saale, 

Germany, e-mail: stoeck@zoologie.uni-halle.de. ' Charles University, Department of Zoology, ViniCnd 7, 

128 44 Praha, Czech Republic. * Universitat Wiirzburg, Institutfiir Humangenetik, Biozentrum, Am Hubland, 

D - 97074 Wiirzburg, Germany. 

Abstract.- A summary of the distribution of green toads containing most of the data published after the discovery 
of polyploid forms (1976) including a map, an index and a bibliography are presented and discussed. 21 Asian 
type localities of hitherto described nominal green toad taxa are shown. The tetraploids are distributed in high 
mountains and extremely continental regions with strong climatic shifts. Records of triploid specimens are 
situated in supposed contact zones between the parapatric diploid and tetraploid toads at foothills of Middle 
Asian high mountains, and triploid bisexual populations occur in the Karakoram and West-Himalayas. Habitats 
of diploids appear to be restricted to lowlands and valley grounds. Polyploids seem to be more resistant. The 
methods hitherto used for the determination of the ploidy level and their applicability are evaluated. We show 
new data on this species complex from Iran including cytometric, karyological, bioacoustic and morphological 
data and we draw taxonomic conclusions for tetraploid Bufo oblongus, diploid Bufo viridis kermanensis, and 
probably diploid Bufo kavirensis. New information on the distribution of triploids in northwestern Pakistan based 
on flow cytometric measurements is presented. The ploidy level of Bufo latastii is revealed to be diploid. 

Key words.- Amphibia, Bufonidae, Bufo viridis complex, Bufo oblongus lectotype, Bufo viridis kermanensis, 
Bufo kavirensis, Bufo latastii, Bufo pseudoraddei pseudoraddei, Bufo pseudoraddei baturae, distribution, type 
localities, Asia, Iran, Pakistan, chromosomes, ploidy determination, calls, systematics 



This paper is a chapter of the doctoral dissertation of M. Stock. 

Introduction 

Since the discovery of tetraploid forms of the Bufo 
viridis complex in the northern Tien Shan (Bachmann 
et al„ 1978; Mazik et al., 1976) many records of dip- 
loid, triploid and tetraploid green toads have been 
made, especially on the territory of the former Soviet 
Union and some few in other countries. New findings 
as the detection of possibly all-triploid, gonochoristic 
populations of this complex in the Karakorum range of 
Pakistan (Stock et al., 1998, 1999) provide arguments 
that the number of investigations in this species com- 
plex will increase during the next years because not 
only zoogeographic and phylogenetic but also cytoge- 
netic, bioacoustic and biochemical questions will be 
studied. Since a fundamental summary of the data on 
the territory of the former Soviet Union was done by 
Borkin et al. (1986a), the development of knowledge in 
this species complex has been dramatically increased. 



Hence, the present paper was aimed to summarize the 
recent information about the distribution which is one 
of the preconditions for the understanding of the natu- 
ral history of these toads. The data on chorology will 
also be an important prerequisite for a revision of sys- 
tematics and nomenclature in this species complex. 

We discuss the hitherto applied techniques of 
ploidy determination, the size of the (known) range of 
polyploid green toads, any detectable correlation of 
their occurrence with climatic/ecological factors, and 
we try to enlarge the knowledge about the ploidy level 
of taxa hitherto described from various type localities. 

Because the knowledge on the ploidy of green 
toads from the territory of Iran and Pakistan is very 
scarce, we also present and discuss some new, sporad- 
ically collected data from different sources, and first 
results of a field excursion to Pakistan. 



Vol. 9, p. 78 



Asiatic Herpetological Research 



2001 




Figure 1 . Map of Middle and Central Asia and parts of the Middle East with records of diploid, triploid and tetraploid 
green toads after 1976. For code numbers see appendix. 



2001 



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Vol. 9, p. 79 











A *loi 








A 


a&£.3^ A 










'^^AA 




□ 




A ^jS A 1 




■Is 






,A 


m 


D bonoaa-e-r\avus (aipioia; 


i 


r 


A Kashgar (tetraploid) 






■ Kapkan (diploid) 



27.5 

I 25 

» 

E 22.5 

o 

1 20 

B 

S 17.5 

« 15 

c 

9 12.5 

co 

«> 10 

3 
<D 

o 7.5 

z 100 150 200 250 300 350 400 

Integrated optical density (Feulgen dye intensity, 
DNA content) 

Figure 2. Diagram showing the nucleus projection area 
in relationship to the integrated optical density (the 
DNA content, respectively) of 100 Feulgen stained 
erythrocyte nuclei of a diploid standard specimen from 
50 km E Gonbad-e-Kavus (3 in Fig. 1), a tetraploid 
standard specimen from Kashgar (43) and the diploid 
specimen from Kapkan (12a). 

Material and Methods 

New data from Iran 

We used three sources: Martens (unpubl.) recorded 
mating calls of green toads from Fasham (4 July 1978, 
23.00) and Polur (30 June 1978, 22.30) which were 
analyzed according to Stock (1998a). Frynta collected 
in 1997 and 1998 (comp. Frynta et al, 1997) green 
toads, among them five living specimens from Ghola- 
man (MTKD D 41350), Kapkan (MTKD D 41351), 
Baghestan (CUP AMPH/IRA/130) and Choqua Zan- 
bil (MTKD D 41352, 41353) whose ploidy levels 
were analyzed with erythrocyte measurements, 
microdensitometrical DNA-measurements and/or 
karyological techniques (Schmid, 1978; Stock and 
Grosse, 1997a). Stock visited in June 1998 Kerman, 
the type locality of Bufo viridis kermanensis Eiselt & 
Schmidtler, 1971, and Birjand, the type locality of 
Bufo oblongus Nikolsky, 1896. A mating call of a sin- 
gle male from Birjand recorded by Stock in the late 
phase of the breeding period (8 June 1998, 21.30) was 
analyzed as described by Stock (1998a). Seven speci- 
mens form Kerman (among them ZFMK 69909 to 
69911, MTKD D 40730, 40731, 41348, 41349) and 
five adult and six juvenile specimens from Birjand 
(among them ZFMK 69901 to 69908, MTKD D 
40729, 41346, 41347) were examined either by flow 
cytometry as described by Stock et al. (1999) or kary- 
ologically according to Schmid (1978). For morpho- 
logical comparisons we included type material - Bufo 
kavirensis: GNM Ba. ex. 1278 (holotype), GNM Ba. 



ex. 1280 (paratype); B. luristanicus: ZMUC R 13221 
(holotype); B. oblongus: ZISP 1952.1 (now lecto- 
type), ZISP 1952.2 (now paralectotype). Institutional 
abbreviations are as listed in Leviton et al. (1985); for 
localities see Fig. 1. 

New data from Pakistan 

In June and July 2000. Stock and Dressel traveled in 
the Northern Areas and North West Frontier Province 
of Pakistan. In the Hunza valley of the Karakoram 
Range near Karimabad (n = 8), and Pasu (n = 50), at 
the tributaries of the Gilgit river near Gupis (n = 2), at 
the Shandur pass (n = 8), and in the Chitral valley, 
near Buni (n = 2) and in Chitral City (n = 8), blood 
samples for ploidy determination of anesthetized 
adult green toads were taken, stored in 70% ethanol 
and refrigerated until flow cytometry according to 
Stock etal. (1999). 

In Skardu, at the western margin of the type region 
("Ladak") of Bufo latastii Boulenger, 1882, blood 
samples of 15 adult toads of this species were taken, 
among them ZMB 62721 to 62726. A mating call of a 
single male (24 June 2000, 21.30) was recorded and 
analysed as described (Stock 1998a). 

For morphological comparisons we examined the 
lectotype of Bufo latastii BMNH 1947.2.21.28 (for- 
merly 72.4.17.223; Stock et al., 1999: Fig. 1). 

Map (Fig. 1), list of records with ploidy detec- 
tion and type localities of nominal Asian 
green toad taxa (Appendix) 

The map contains most of the localities or regions 
where diploid, tetraploid and/or triploid toads have 
been recorded in Middle and Central Asia and the 
eastern parts of the Middle East after 1976, the year of 
the first detection of polyploids. Our map covers all 
regions where polyploid green toads have been found. 
These records (numbers 1 to 74) from the literature 
have been made using various methods for identifica- 
tion or determination of the taxa and/or ploidy levels. 
Some methods are not unambiguous and might there- 
fore cause errors. Consequently, it was necessary to 
evaluate the methods which were utilized for the 
determination of the ploidy level because not all data 
are indisputable. In the case of the numbers 1 to 74, 
the present article only includes papers if they either 
contain statements on the ploidy of the toads or data 
appearing suitable to draw a conclusion to the ploidy 
(e.g. call data, see below). However, even if authors 
distinguished between diploid and tetraploid toads, in 
rare cases, especially in abstracts, the localities of the 
records were not precisely published (e.g., Fikhtman, 
1989) or it was impossible to deduce from the publi- 



Vol. 9, p. 80 



Asiatic Herpetological Research 



2001 



20 
175 

IS 
12.5 

10 













A 










* A 








■ 




*.*4t£\4 


M* 






■ 
■ 


* * 4 d 


SSftf*^* 4 






rA-i" 


a J»*~i 






i^Hrv i 








D™ 


TT 




A Kashgar (tetraploid) 


D 






H Gholaman (diploid) 
1 i 



50 100 150 200 250 300 350 

Integrated optical density (Feulgen dye intensrtsy; DNA-content) 



Figure 3. Diagram showing the nucleus projection area 
in relationship to the integrated optical density (the 
DNA content, respectively) of 100 Feulgen stained 
erythrocyte nuclei of a diploid standard specimen from 
S of Gorgan (2), 50 nuclei of a tetraploid standard 
specimen from Kashgar (43), 100 nuclei of the tetrap- 
loid specimen from Baghestan (13) and 100 nuclei of 
the diploid specimen from Gholaman (1). 

cation to the exact ploidy of toads from a concrete 
locality (e.g., Castellano and Giacoma, 1998) and/or 
the method of ploidy determination was not men- 
tioned (e.g., Pisanets & Vasilenko, 1995). Since no 
polyploid green toads have been found in the Cauca- 
sus (e.g., Kuzmin 1995: 187), we disregarded this 
area. If the information about a region but not a spe- 
cial position was available, the record is not shown in 
the map but listed in the appendix, and marked with a 
"?" (instead a number) in the line next to the nearest 
concrete record. Furthermore, we present some hith- 
erto unpublished single records ("Stock, unpubl.") 
from Kazakhstan and Kyrgyzstan. 

We used different maps (Anonymous, 1993; 
DMAAC) and/or the descriptions and sketch maps in 
the literature (e.g., Borkin et al., 1986a) for the local- 
ization of the records. If they were available, the 
appendix contains the condensed descriptions of the 
localities, the geographic position and the method of 
ploidy determination. 

The second part of the appendix (letters A to V) 
contains the 21 Asian type localities shown in Fig. 1 
and represents a preliminary precondition for a sys- 
tematic discussion. The bibliography in that part only 
comprises a choice of the literature which either dis- 
cussed the systematic rank of a taxon or shows details 
suitable to draw a conclusion to the ploidy. Some 
descriptions of taxa either mentioned regions only but 
no special type locality (e.g., Boulenger, 1882: 
"Ladak" for B. latastii), but were drawn in the map; 
others presented a confusing diversity of type locali- 



01 

E 

3 



o 




iiriiiiliifftinl! 

20 40 60 



80 100 120 140 160 180 200 



DAPI-fluorescence 
Figure 4. Histogram obtained by DNA flow cytometry 
from a mixture of DAPI stained blood samples with 
chicken as the standard (a), CV = 2.63%, a diploid 
green toad from Kerman, type locality of Bufo viridis 
kermanensis, (b), CV = 3.62%, and a tetraploid green 
toad from Birjand, type locality of Bufo ob/ongus, (c), 
CV = 3.31%. Total cell number 10 327. 

ties covering large parts of Central Asia (e.g., Bed- 
riaga, 1898: B. viridis var. pewzowi and var. strauchi 
with a type series from 4 or 14 localities, respectively, 
distributed from Mongolia to the Pamirs). In the latter 
cases, only some localities are shown, and this is men- 
tioned in the appendix. In future, such taxonomic 
problems should be resolved by careful lectotype des- 
ignations leading to type locality restrictions as a 
basis of a revision. Finally, the status of some old 
names, at least for diploid green toads, still remains 
unclear (see ref. in Kuzmin, 1999: 251, 264). They 
were not shown in the map or originated from regions 
outside of it. 

Results and Discussion 

New data from Iran 

Cytometric and karyological data. A large male 
from Kapkan (Fig. 1: 12a) in the East-Iranian part of 
the Kopet Dagh which we considered to represent 
Bufo viridis turanensis was identified to be diploid 
(Fig. 2). A male Bufo viridis ssp. from Gholaman 
(Fig. 1:1; Fig. 8C) in the Zagros mountains, was dip- 
loid (Fig. 3); two additional males from Choqa Zanbil 
(Fig. 1: la) below the south-western foot of the 
Zagros mountains were also diploid. 

All toads examined from Kerman, the type local- 
ity of Bufo viridis kermanensis, were found to be dip- 
loid as well (Fig. 4). This agrees with the 
contemporaneously published results of Borkin et al. 
(2000). We detected (Fig. 3) the first tetraploid toad in 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 81 





c 


!H 


- — 


•ill*- 


ins ii b 

12 3 4 

•1 W XX AA 

5 6 7 8 

IX III M 

9 10 11 


Ii it ii a 

12 3 4 

II 11 |« M 

5 6 7 8 

* * «■ ->. 

9 10 11 



Figure 5. Chromosomes of diploid male Bufo viridis 
kermanensis Uom Kerman. a - Giemsa-stained 
metaphase, b - Giemsa-stained karyotype, c - Q- 
banded metaphase, d - Q-banded karyotype. 

Iran near Baghestan (Fig. 1: 13; Fig. 8F) in the moun- 
tainous region north-eastern of the Central Iranian 
Plateau (abstract by Stock et al.. 1998). A sample of 
five toads from Birjand (Fig. 1: 13a), the type locality 
(Fig. 1: E) of Bufo oblongus, was found to be tetrap- 
loid (Fig. 4). Table I outlines the results of the flow 
cytometric measurements in specimens from Kerman 
and Birjand. Table 2 summarizes the results of eryth- 
rocyte size measurements. The erythrocyte sizes were 
typical of diploid and tetraploid green toads, respec- 
tively, according to Stock and Grosse (1997a). The 
mean DNA content of B. oblongus (17.02 pg) rather 
corresponds to that of tetraploids from Kashgar (17.5 
pg; Stock, 1998b) measured with the same technique, 
but during another session. 

The Giemsa-staining (Fig. 5a, b) revealed a diploid 
karyotype of B. viridis kermanensis. These convention- 
ally stained chromosomes posses the characters of 
those from some other Eurasian diploid Bufo viridis 
which have been examined until now (Birstein. 1981; 
Bogart. 1972; Roth and Rab. 1987; Schmid. 1978: 
Ullerich. 1966). The pair 6 terminally exhibits in its 
long arms a secondary constriction which is caused by 
the telomeric position of the nucleolus organizer 
regions (NORs) in B. viridis (see also Roth and Rab, 
1987; Schmid 1978). The Quinacrine-banding in 
chromosomes of B. viridis kermanensis (Fig. 5c. d) 
shows differently intensive fluorescing chromosomal 
regions in pairs 6 to II, but distinct bright fluorescence 



»» . 

1 h m 

1 2 3 

Mi mi in* 

4 5 6 

XXXX MAI MM 

7 8 9 




1 * 
UJVMM 

4 5 6 

■vkX «MI xxk» 



Figure 6. Chromosomes of a tetraploid female Bufo 
oblongus Uom Birjand. a - Giemsa-stained metaphase, 
b - Giemsa-stained karyotype, c - Q-banded 
metaphase, d - Q-banded karyotype. 

was only found in short and long arms of pairs 6 and 8. 
Birstein (1981) detected Q-bands in a B. viridis from 
the Crimea in pairs 6 to II (the most distinct ones in 
both arms of pairs 7, 10, and II). In a triploid taxon 
(Bufo pseudoraddei balurae) of the B. viridis complex 
from the Karakoram. we found Q-bands in the long 
arms close to the centromere of triplet 1, in the short 
arms of triplets 6 and 7 and in both arms of triplets 8 to 
11 (Stock etal., 1999). 

Similarly, the karyotype of the tetraploid Bufo 
oblongus exhibits Q-bands in chromosomes of the 
quartets 1, 6 to 11 (Fig. 6c. d). The most interesting 
linding is the occurrence of Q-bands in the short arms 
of only two out of the chromosomes of quartet 6. These 
two chromosomes have also larger long arms than the 
two remaining Q-negative chromosomes, and there- 
fore, we conclude that the Q-positive pair also repre- 
sents the only one which carries the telomeric NORs as 
Roth and Rab (1987) found in tetraploid toads from 
Kyrgyzstan. We are preparing a detailed cytogenetic 
study. Like in the triplet 1 of triploids from Karakoram. 
one or two chromosomes of quartet I in B. oblongus 
posses Q-bands in their long arms. Furthermore, addi- 
tional differences are visible in the occurrence and 
position of Q-bands among the chromosomes in each 
of the quartets 7. 8 and 10 (Fig. 6c. d). These observa- 
tions in B. oblongus provide arguments for considering 
this tetraploid form allopolyploid. The question of 
autoploidy or alloploidy of polyploids from various 



Vol. 9. p. 82 



Asiatic Herpetological Research 



2001 



Table 1 . DNA content in diploid toads from Kerman (Bufo viridis kermanensis), diploid toads from Choqua Zanbil 
(B. wridisssp.), tetraploid toads from Birjand [Bufo ob/ongus), and diploid B. iatastiiUom Skardu measured by flow 
cytometry of DAPI stained erythrocytes with chicken nuclei (= 2.34 pg DNA/nucleus) as a standard. SD = standard 
deviation. 



Kerman, diploid 


C. 


Zanbil, 


diploid 


Bi 


rjand, tetraploid 


Skardu, 


diploid 




(N = 7) 






(N = 


2) 






(N = 


5) 






(N = 


15) 



















. . 








. , 








, ~ 






*~ 








1 ~ 








* , ~ 








ii 




f . 


ii 






^^ 


ii 






, , 


n 






^^ 


s 




en 
Q. 


■o 






CO 
Q. 


■o 






Q. 


"D 






D. 


co 




— 


CO 








CO 








CO 








■d 

c 

3 




C 
0) 


c 

CO 






C 
CD 


CO 






C 
CD 


■o 

c 

CO 






c 

CD 
































s? 


o 




5? 




O 




s? 




O 




S? 




O 


c 




o 


c 






O 


c 






CJ 


c 






O 


2 


c 


< 


<D 


c 




< 


CD 


c 




< 


CD 


c 




< 


o 


CO 


z 


o 


CO 




Z 


O 






Z 


CJ 






Z 


o 


CD 

5 


Q 


o 


^ 




Q 




2 




Q 
CD 


.c 

CJ 


5 




Q 

CD 


o 


o 




o 


o 






o 


o 






o 


o 




































o 


o 


o 


o 


o 




o 


o 


o 




O 


o 


o 




o 














CO 








CO 








CO 


co 


CO 


n 


CO 


CO 




.O 


CO 


CO 




n 


CO 


CO 




.O 


cr 


cr 


< 


CL 


cr 




< 


CC 


H 




< 


cr 


cr 




< 


Mean 3.43 


100 


8.02 


3.59 


100 




8.42 


7.27 


100 




17.02 


4.13 


100 




9.68 



Mm 3.25 94.75 7 61 3.56 99.04 8.34 7 04 96.83 16.47 3.77 91.21 8.83 

Max 3.69 107.58 8.63 3.63 100.95 8.50 7.57 104.13 17.71 4.59 1 10.95 10 74 

SD 0.17 - 40 0.05 - 0.11 0.19 - 0.45 0.25 - 0.27 



Table 2. Erythrocyte size (projection areas of 30 red blood cells per toad were measured) in five tetraploid speci- 
mens from Birjand (B. ob/ongus), a tetraploid specimen from Baghestan, six diploid specimens from Kerman (B. 
viridis kermanensis), a diploid specimen from Kapkan, a diploid specimen from Gholaman, and two diploid speci- 
mens from Choqua Zanbil. 



Parameter (urn 2 ) 



Locality 
(ploidy) 



ii 



CO M 

iS 



-5, " 
01 r- 
S 5 



CO 

ii 



!o N 

1" 

HI C flj c 



C CM 

CO <N 
E M 



II 

C Z 
(0 — 



CO 



C\l 



Mean of means in the population 

Maximal mean in the population 

Minimal mean in the population 

Largest cell measured 

Smallest cell measured 

Mean standard deviation in the pop. 



353.87 
374.30 
333.03 
463.29 
274.07 
33.62 



335.98 
335.98 
335.98 
413.50 
280.11 
30.80 



256.38 
274.93 
224.40 
358.91 
183.95 
23.22 



272.24 
272.24 
272.24 
318.11 
216.37 
21.99 



248.39 
248.39 
248.39 
322.77 
216.21 
22.17 



CN 

II 

z 



O CL o 21 



236 26 
181.47 
286 32 
181.47 
286.32 
22.94 



localities has been controversially discussed (for over- 
view see Balletto et al., 1999: Stock et al.. 1999). As 
compared with conventionally stained tetraploid karyo- 
types (e.g.. Borkin et al.. 1986b, c: Borkin and Kuzmin. 
1988; Orlova & Uteshev, 1986; Pisanets. 1978; Roth & 
Rab, 1986, 1987; Stock. 1998b; Toktosunov, 1984; 
Whu & Zhao. 1987). that of B. oblongus (Fig. 6a. b) 
does not exhibit visible differences. 



Mating calls (Fig. 7, Table 3). The mating call data 
from Lar valley (Andren and Nilson, 1979), Fasham 
and Polur suggest that these toads are probably diploid 
(see below for methodical questions). The same predic- 
tion (Stock. 1998a) is possible for the toads from 
Cheshmeh-ye-Sefid-Ab, the type locality of Bufo kavi- 
rensis. The mating call data from Polur (2a) and 
Fasham (2b) and the data from Lar valley (2b) and 
Gorgan (2) refer to the occurrence of (only) diploid 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 83 



35 



30 



n 25 

<D 
*■» 

n 

a> 20 
v> 

3 

a. 



15 



10 









/ 


<► 
<> 






,- 




<> 
<> 






/ 








, * 


' 


A" 

A 

A - 

'A 




r 




■■ 
■■ 

H 

* 








1 

1 «.^i 





s 






M 


A M 











A 
▲ 
D 



Tetraploid. Middle and Central Asia 
(Stock, 1998a, b) 

Birjand 

Diploid, Middle Asia (Stock, 1998a) 

Lar Valley (Andren and Nilson, 
1979) 

Cheshmeh-ye-Sefied-Ab (Andren 
and Nilson, 1979) 

Fasham 

Polur 

Linear (Tetraploid, Middle and 
Central Asia (Stock, 1998a. b)) 

• Linear (Diploid, Middle Asia (Stock, 
1998a)) 



10 15 20 25 

Water temperature (°C) 



30 



Figure 7. Pulse rate of mating calls of green toads from Iran in comparison with pulse rate of diploid and tetraploid 
toads from Middle and Central Asia. Birjand: type locality of Bufo oblongus Nikolsky, 1 896. Toads from Birjand 
were found to be tetraploid (Fig. 4). Cheshmeh-ye-Sefid-Ab: type locality of Bufo kavirensis, Andren and Nilson, 
1979: the call data appear to elucidate that the taxon is diploid. Call data from Lar valley (Andren and Nilson, 
1979), Fasham and Polur (present study) provide arguments that these toads are diploid. 



green toads in the Elburz mountains and confirm the 
evolutionary conservatism of the mating calls (disc, in 
Stock, 1998b; Stock et al., 1999). The single mating 
call of a tetraploid male ( 1 3a) from the type locality, 
Birjand (E), of Bufo oblongus confirmed previous data 
on tetraploids from various regions of Middle and Cen- 
tral Asia (Castellano et al., 1998; Stock, 1998a, b). 

Morphology and Taxonomy (Fig. 8). Although the 
locality (1) is relatively close to the type locality (A) 
of Bufo luristanicus (Schmidt, 1952), the morphology 
of the toads from Gholaman (Fig. 8C) and Choqa 
Zanbil differs completely from that of the B. luristani- 
cus holotype (Fig. 8H). This confirmed once more the 
occurrence of at least two different green toad taxa in 
the north-western Zagros mountains of Iran as already 
Schmidtler and Schmidtler (1969) as well as Eiselt 
and Schmidtler (1973) stated. First, the form which 
was previously (Schmidtler and Schmidtler, 1969; 
Mertens, 1971b; Eiselt and Schmidtler, 1973) called 
B. viridis arabicus and to which the diploid toads 
from Gholaman and Choqa Zanbil belong to. Now, 
the name "arabicus" is no longer applicable since Bal- 
letto et al. (1985) restricted it to B. arabicus Heyden, 
1827 from the Arabian Peninsula. Therefore, we pre- 
liminarily use the term Bufo viridis ssp. for them. The 
second taxon occurring in the north-western Zagros 



mountains until eastern Iraq (Afrasiab and Ali, 1988) 
is B. (surdus) luristanicus. As to be seen in the appen- 
dix, the ploidy level of all three subspecies of B. sur- 
dus distinguished by Schmidtler and Schmidtler 
(1969) and Eiselt and Schmidtler (1973) is still 
unknown (cp. Baloutch and Kami, 1995). 

The seven topotypic specimens of Bufo viridis 
kermanensis from Kerman (13b) are very similar to 
the holotype (NMW 19647) shown and described by 
Eiselt & Schmidtler (1971, 1973). This diploid form 
is different from the large sized diploid B. viridis 
turanensis as already Hemmer et al. (1978) stated and 
which was confirmed by our data. 

The diagnostic differences between Bufo kaviren- 
sis and B. viridis kermanensis consist according to 
Andren and Nilson (1979) in only two morphometric 
traits (I: ratio "distance between posterior border of 
nostril and anterior corner of eye/internasal distance" 
0.80 - 1.06 in B. kavirensis and 1.21-1.55 in B. v. ker- 
manensis; II: ratio "width of upper eyelid /interorbital 
distance" 1.1 1 - 1.61 in B. kavirensis and 1.51 - 2.00 
in B. v. kermanensis). Both ratios were not confirmed 
because the first ranged for our seven B. viridis ker- 
manensis from 0.9 - 1.18 (and 0.93 - 1 .05 in two spec- 
imens, Fig. 1: 13c, by Borkin et al., 2000); the second 
ratio varied also strongly from 0.8 - 2.92 (and 1.83 - 



Vol. 9, p. 84 



Asiatic Herpetological Research 



2001 



2.31, Borkin et al., 2000). Including the data shown 
above, we preliminarily consider (diploid) B. kaviren- 
sis to be a junior synonym of diploid B. viridis ker- 
manensis. This also agrees with the range presumed 
by Eiselt and Schmidtler (1973) for their taxon. The 
relationships between diploid B. viridis kermanensis 
and diploid B. viridis ssp. (previously called B. viridis 
arabicus, see above) require additional investigations. 

The morphological differences between Bufo 
oblongus (Fig. 8A, B, D) and B. viridis kermanensis 
(Fig. 8E) which were considered to be "very sharp 



regarding neighboring races" (Eiselt and Schmidtler. 
1973) were now explained by the different ploidy lev- 
els. The females of B. oblongus (e.g., Fig. 8B) exam- 
ined from Birjand exhibited a coloration pattern 
which is very similar to that of the specimen ZISP 
1952.1 from the type series (Fig. 8A). We therefore 
designate it here as the lectotype of B. oblongus, 
Nikolsky, 1896, and we consider this taxon according 
to the topotypes to be tetraploid. This demonstration 
of tetraploidy for B. oblongus is important for the tax- 
onomy of green toads in Asia because the name is 



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2001 



Asiatic Herpetological Research 



Vol. 9. p. 85 



Table 4. Mating call data of triploid Bufo pseudoraddei baturae'xn comparison with Bufo /atasti/Uom India and 
Pakistan. 





























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S 








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B. lalastii 


Kargil and Tang- 


no 


no 


30- 


8-13 


1200-1300 


2.5-6.1 


no data 


Dubois and 




marg, 


Pahalgam 


data 


data 


60 






[data from 
spectrograms] 




Martens 
(1977) 


B. lalaslii 


Skard 


u(49) 


13 


54.3 


39.8 


10.62 


1470 


4.3-6.9 [3] 


4.9-69 


present 


(diploid) 


















[3] 


paper 


B. pseudorad- 


Gilgit (48a) and 


7-27 


27.3- 


19- 


7.38- 


1378-1636 


1.2-4.2 


2.0-6.8 


Stock et al. 


dei baturae 


Pasu (47, P) 




86.2 


56.7 


21.28 




[25] 


[25] 


(1999). 


(triploid) 




















unpubl. 



(one of) the oldest available name(s) for tetraploids in 
Middle and Central Asia. 

New data from Pakistan 

Cytometric data. Among the triploid populations 
reported by Stock et al. (1999), DNA-measurements 
in additional specimens confirmed bisexual triploidy 
in altogether 82 adult specimens of Bufo pseudorad- 
dei baturae from Pasu (47). Exclusively triploids were 
also detected by our sample survey along the upper 
tributaries of the Hunza (47a) and Gilgit river systems 
(48b, c). The record of triploids at the Shandur pass 
(48c, 3720 m a.s.l ) and the occurrence of triploids in 
the Chitral valley (48d, e) demonstrated their continu- 
ous distribution, i.e. a more or less connected range. 
In Chitral, sympatric occurrence of triploids with B. 
stomaticus was observed. 

The flow cytometric measurements (Tab. 1, Fig. 
9E) revealed all 15 specimens of Bufo latastii from 
Skardu (49) to be diploid. Although the absolute val- 
ues cover a relatively large range resulting from the 
fixation of samples in ethanol in the field, apparently 
the DNA content of B. latastii is about 15 to 20% 
higher than that of diploid B. viridis ssp. and B. v. ker- 
manensis from Iran. The karyotype of toads from 
Skardu comprises 2n = 22 chromosomes; a detailed 
cytogenetic study will be published elsewhere. It 
remains to be clarified if the karyological study on a 
"Bufo spec." from Kashmir (Duda & Opendar, 1971) 
really represented B. latastii as Dubois and Martens 
(1977) and Roth and Rab (1986) presumed when they 
considered this species diploid. In any case, long last- 
ing speculations (see appendix: M) on the ploidy of B. 
latastii which previously were connected with signifi- 



cant uncertainties in the systematics of Central Asian 
green toads (Borkin et al. 2000; Stock et al., 1999) 
have been finished now. 

Mating calls (Table 4). Mating call parameters of a 
single diploid male B. latastii from Skardu were 
found in the range reported for this species by Dubois 
and Martens (1977) from Kashmir and Ladakh. As 
discussed by Stock et al. (1999), the B. latastii calls 
are rather similar to those of triploid B. pseudoraddei 
baturae from Western Karakoram and B. p. pseudo- 
raddei from Western Himalayas. This can be either 
interpreted as convergences in isolated species or may 
result from the participation of B. latastii genes in the 
probably allotriploid B. p. baturae, but up to now, 
exact explanations are not available. 

Morphology and taxonomy. The morphology of 
triploids from Shandur Pass (48c) and Chitral (48d, e) 
was similar to that of B. pseudoraddei baturae from 
Gilgit (Stock et al., 1999); therefore, we consider 
them to represent the same taxon. 

The 15 toads examined from Skardu (49) were 
clearly identified as Bufo latastii Boulenger, 1882 (= 
B. siachinensis Khan, 1997). The external morphol- 
ogy and the coloration are very similar to the lecto- 
type of B. latastii (Fig. 9A-D). Furthermore, the 
characteristic ventral dark pattern comprising black- 
ish, up to eye-sized, sometimes connected spots and 
an abrupt darkening of the caudal third of the belly 
including the base of the hind legs was found to occur 
in the B. latastii lectotype, the B. siachinensis holo- 
type, and always in the toads from Skardu also (Fig. 
9C, D). 



Vol. 9, p. 86 



Asiatic Herpetological Research 



2001 



Table 5. Evaluation (overview) of methods previously used by different authors for the determination of diploid and 
polyploid green toads. For detailed information see text. 



Method of determination 



Unequivocalness 



Applicability in the field Abbr. in the appendix 



Karyotype, karyogram, counting of 
chromosomes 

Flow cytometry 



Microdensitometry (Feulgen stain- 
ing), cytometry 



Electrophoresis 
Measurement of erythrocytes 



yes 

yes, if applied professionally 

yes, if applied professionally 

no, only prognosis possible 
no, only prognosis possible 



probably yes, for distinguish- 
ing between diploid and poly- 
ploid (triploid. tetraploid) 
forms, but see B. latastii 



External morphology, morphometry no, only prognosis possible 
Form and size of the clutch no, often misleading 



Mating call analysis 



no 



no, but blood samples can 
shortly be stored in etha- 
nol 

yes, if blood smears are 
stored in 5% formaldehyde 
solution 



collection of air dried blood 
samples easily possible 

yes 



yes 



chrom. count, 
flow cytom. 

microdens., cytom. 

electrophor. 
erythr. size 

calls 



ext. morph. 
clutch 



Evaluation of methods for the identification 
and determination of the ploidy in Asian 
green toads (Table 5) 

Some forms/species of Middle and Central Asian 
green toads have been partly identified using mislead- 
ing or ambiguous methods (comp. also Stock and 
Grosse, 1997a). 

The karyological analysis still remains an essen- 
tial tool. The treatment with colchicine and the prepa- 
ration of bone marrow, spleen or intestine tissue for 
the counting of stained chromosomes (e.g., Schmid, 
1978; Macgregor and Varley, 1983) requires the kill- 
ing of the animals. The obtaining of blood by heart 
puncture with micro-syringes (e.g., Schroer, 1996) 
and the determination of the ploidy by chromosome 
counts from blood cultures (Castellano and Giacoma, 
1998) causes little mortality but requires the transport 
of the toads to the laboratory. 

Flow cytometry is another unequivocal technique 
(Murphy et al., 1997 for review). The preferable stor- 
age of blood by freezing in liquid nitrogen is usually 
limited in the field. Ethanol fixation of blood obtained 
from anaesthized toads allows a storage for some 
weeks and the ploidy determination by flow cytome- 
try if refrigeration is possible (e.g.. Stock et al., 1999), 
but does not allow the exact determination of absolute 
DNA-contents. 

Feulgen staining of erythrocytes and the 
microdensitometrical determination of their DNA- 



content provides also unequivocal results (Stock and 
Grosse, 1997a). Although better results can be obtained 
if applied in the laboratory since the storage of air dried 
blood causes a loss of stainability, we recently got good 
results with blood smears which were stored in 5% 
formaldehyde solution until staining (our unpublished 
data). The obtaining of blood smears from the tip of the 
finger seems to be of very little damage for the animals, 
but requires to prevent the pollution of blood samples 
with skin mucus. 

Significant differences in the average erythrocyte 
size allow a prognosis of the ploidy level (Stock and 
Grosse 1997a). Discrimination problems occur 
because some populations exhibit values in the over- 
lapping range between diploid and tetraploid toads. In 
addition, the classification of triploid individuals 
remains unsolved. In probably all-triploid populations 
from Karakoram, the mean erythrocyte areas exhib- 
ited significant differences to diploid and tetraploid 
populations (Stock et al., 1999), but the variability of 
the individual erythrocyte size shows a large overlap- 
ping with diploid and tetraploid specimens. Unfortu- 
nately, these phenomena also prevent an exact 
analysis of the ploidy level of museum and type spec- 
imens with the technique of Mercadal (1981). 

Results of protein electrophoresis (e.g., Borkin 
and Sokolova 1989, Mezhzherin and Pisanets, 1995a, 
1995b) can usually not substitute the preceding ploidy 
determination since "electromorphs" may cause mis- 
interpretations. Some authors of electrophoretic stud- 



2001 



Asiatic Herpetological Research 



Vol. 9. p. 87 




Figure 8. A - Lectotype (present designation) of Bufo ob/ongusN\ko\sky, 1896 (Z ISP 1952.1) from Birjand, E-lran. 
B - Topotypic tetraploid female Bufo oblongus\xom Birjand. C - Diploid male from Gholaman. D - Topotypic tetrap- 
loid male Bufo ob/ongusUom Birjand. E - Topotypic diploid male of Bufo viridis kermanensis Eiselt & Schmidtler, 
1971. F - Tetraploid male from Baghestan. G - Holotype of Bufo kavirensis, Andren & Nilson. 1979. H - Holotype of 
Bufo luristanicus Schmidt, 1952. 



Vol. 9, p. 88 



Asiatic Herpetological Research 



2001 



ies did not describe whether or how they determined 
the ploidy level (e.g.. Mezhzherin and Pisanets. 
1991). 

At any given temperature mating calls of tetrap- 
loid toads exhibited longer pulses and interpulse inter- 
vals resulting in lower pulse rates than diploid toads 
(Stock 1997a. 1998a). a finding which exhibits paral- 
lels in other diploid/tetraploid anuran species (Hyl- 
idae, Leptodactylidae) and artificial tetraploids 
suggesting that differences are caused by polyploidy 
(Gerhardt. 1994: 317). 

For triploid toads from northern Kyrgyzstan (Cas- 
tellano et al.. 1998) and triploid populations from 
northern Pakistan (Stock et al.. 1998. 1999) call 
parameters were found to resemble those of tetraploid 
toads. Therefore, the mating call analysis seems suit- 
able for distinguishing between diploid and polyploid 
(triploid. tetraploid) green toads. However, indepen- 
dently from polyploidisation. speciation in the B. viri- 
dis complex appears to take place also within each 
ploidy level and as in other anurans. evolutionary 
divergences in sympatry as well as convergences of 
call parameters in allopatry can be expected. The lat- 
ter seems to be the case in diploid B. latastii. from 
which pulse rate data (Dubois and Martens. 1977) 
suggested a polyploid species (Stock, 1998a). 

Many morphometric traits exhibited differences of 
the means but values showed large intersections 
between both diploid and tetraploid toads and mor- 
phometric parameters were not suitable for ploidy 
determination (Stock. 1997b). The study announced 
by Roth (1986) on "about 2000 specimens from the 
whole range" has never been published (Roth. pers. 
comm.). In advance. Roth (1986) considered morpho- 
logical characters "useless" for taxonomy but without 
determination of the ploidy level of the collection 
material. Multivariate analyses confirmed relation- 
ships between the ploidy level and the external char- 
acter syndrome in diploid and tetraploid green toads 
(Stock, 1997b; Castellano et al.. 1998). Triploid toads 
from Kyrgyzstan exhibited more distinct differences 
to diploid than to tetraploid specimens (Castellano et 
al., 1998). Triploid populations in the Karakoram 
showed significant differences to both diploid and tet- 

Figure 9. A, B - Lectotype of Bufo /<?/<3s///Boulenger, 
1882 (BMNH 1947.2.21.28, formerly 72.4.17.223), 
scale for B: 1 centimetre. C, D- Male B. /atast//(ZMB 
62721 ) examined from Skardu, Baltistan. E - Histogram 
obtained by DNA flow cytometry from a mixture of 
DAPI stained blood samples with chicken as the stan- 
dard (a), CV = 2.94%, and diploid B. /atasti/Uom 
Skardu (b), CV =-2.74%. Total cell number 22 744. 



raploid toads from Central Asia in many characters 
with univariate methods (Stock et al., 1999). A prog- 
nosis for the purely morphometric classification of 
diploid, triploid and tetraploid toads is possible with 
multivariate methods and this appears to be one tool 
for the prognostic assortment of preserved specimens 
in collections. The prognosis can be improved if mor- 
phometric data are combined with erythrocyte size 
data (Stock, 1997b; Stock and Grosse, 1997a; Stock et 
al., 1999). In any case, the use of exact methods for 
ploidy determination is highly recommended (e.g., 
Borkin et al.. 2000). 

Pisanets (1987) indicated differences in form and 
size of the clutch between diploid and tetraploid toads 




\ r— -1 r- 

20 40 60 80 100120140160180200 



DAPI-tluorescence 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 89 



from Turkmenistan (Kurukhaudan village. 12). The 
character is occasionally misleading (Kuzmin, 1995: 
94; Stock and Grosse, 1997a) although it was infre- 
quently used for the identification of toads for further 
investigations (e.g., Mezhzherin and Pisanets, 1990). 

Distribution 

In general, the present map (Fig. I) does not com- 
pletely reflect the occurrence of green toads whose 
ploidy is known but also illustrates the recent degree 
of investigation. 

Diploid toads (black arrows without starlet). The 

records of green toads found to be diploid in Middle 
Asia are distributed in the Turan Basin and are also 
concentrated at the foothills of the high mountains. 
Diploids have been found along the bottom part of the 
Kopet Dagh mountains (7, 8, 10, 1 1); a second group of 
records is to be seen in the Kafirnigan and Vakhsh river 
valleys (23, 25, 28, 29). The last records seem to exem- 
plify a dispersion of diploid toads along the rivers into 
the mountain valleys. The western and particularly the 
northern Tien Shan form a third region with several 
records of diploid toads (31a, 36, 38, 39, 59, 59a). 
More distant from the mountains researches combined 
with ploidy determination have only sporadically been 
carried out. Merely some localities with doubtless dip- 
loid toads near the lower Amu-Darya (9, 14) and Zer- 
avshan river (17, 18) are known. The record of diploid 
toads from the Chingzhal river (61) in Kazakhstan by 
Golubev (1990) appears to be doubtful (Dujsebayeva et 
al., 1997). The main range (maps in Dubois and Mar- 
tens, 1977, Stock et al., 1999) of the diploid B. latastii 
(N, 49) is the rather humid Kashmir valley and the 
surrounding rather arid mountain ranges. 

The small and highly generalized map by Kuzmin 
(1995: 182) and the more detailed one (Kuzmin, 
1999: 479) show either a continuous range or many 
separate symbols of (diploid) B. viridis in Middle Asia 
and Kazakhstan. Both maps cannot reflect the detailed 
records and/or were not based on karyological studies. 
Therefore, Kuzmin (1995, 1999) also announced that 
the distribution requires additional investigations. 
However, Kuzmin's maps refer to the occurrence of 
diploid green toads in the Turan Basin along the rivers 
whereas the symbols for (diploid) B. viridis in the 
Inner Tien Shan and around the Issyk-Kul appear to 
be very doubtful (e.g. see Borkin, 1989). 

In spite of only few references which contained 
data on the ploidy level and on the altitudinal distribu- 
tion, the occurrence of diploid toads in Middle Asia 
apparently exceeds 1600 m a.s.l. only in rare cases 
(appendix; Castellano et al., 1998: "At least in Kaza- 
khstan and Kyrgyzstan no diploid populations have 



been found above 2000 m a.s.l."). Many papers (sum- 
marized e.g. by Meinig, 1995) reporting the occur- 
rence of "B. viridis" until 4000 m a.s.l. in Central 
Asia, very probably represent data on polyploid toads. 
This also may account for Kuzmin's (1999) material 
who reported on B. viridis up to 3200 m a.s.l. in Mid- 
dle Asia. 

Tetraploid toads (white arrows). The tetraploid 
toads are largely distributed in the Middle and Central 
Asian mountain systems. They have been described 
from the Kopet Dagh (5, 6, 8, 11, 12), the Gissar range 
including its southern spurs (21 to 27, 27a, 30), the 
western Tien Shan (31a, 32, 33, 40), the northern Tien 
Shan and the Issyk-Kul gorge (39, 51 to 59, 59b, 60), 
the Central Tien Shan (42, 51), the margins of the Tak- 
lamakan desert (43, 50?), the region between Zailiyskiy 
Alatau and lake Balkhash (60, 37, 37a, 37b), the Dzun- 
garian Alatau (60a, 60b), the eastern Tien Shan (63), 
and Dzungaria (62, 64, 66 to 72). In the Pamirs, tetrap- 
loid toads have been found in the western (34, 35) and 
in the central parts (45). Furthermore, Stock (1998b) 
found toads from eastern Pamirs (44) to be tetraploid. 
They were recently described by Fei et al. ( 1999) as a 
new subspecies (T). The present map provides argu- 
ments for a rather continuous distribution of tetraploid 
toads from Eastern Tien Shan to the Dzungarian Gobi 
in Mongolia. Interestingly, eastern of about 80° E only 
records of tetraploids and obviously no record of dip- 
loid toads have been made. Various records underline 
the high ecological plasticity especially of the tetrap- 
loid green toads (e.g. Dujsebayeva et al., 1997). The 
dry centers of the large continental deserts Kyzylkum, 
Karakum, Muyunkum and Betpak-Dala seem to be 
not suitable as habitats of (tetraploid) green toads but 
this might be also caused by the recent low degree of 
investigation. Considering some records of tetraploids 
in the eastern plane regions, some additional records 
of tetraploid toads in the Turan lowlands can be 
expected in future. The tetraploids in the West of the 
known range (Kopet Dagh) may have a rather contin- 
uous distribution via the mountains of Afghanistan 
until those of the Pamiro-Alai-System, but data from 
Afghanistan are recently too scarce for a detailed 
analysis. 

Triploid toads (black arrows with starlet). For the 

first time, probable triploidy of three specimens was 
detected by Bachmann et al. (1978: "36% more DNA 
than diploid Bufo viridis" ) in toads (presumably) orig- 
inating from Kabul (48f in Fig.l; Hemmer et al., 
1978: 352, 370). Triploid individuals have been 
detected in south-western Turkmenistan (5), in north- 
ern Kyrgyzstan (39) and south-eastern Kazakhstan; 
the last record was reported without particular locality 



Vol. 9, p. 90 



Asiatic Herpetological Research 



2001 



(Borkin et al., 1997, 2000) and we did not draw it in 
the map. The rare occurrence of triploid individuals in 
the Central Pamirs (45) has been concluded from 
isozyme data (Mezhzherin and Pisanets 1990). Most 
of the authors supposed a hybrid origin of triploids as 
a result of mismatings between diploid and tetraploid 
toads or only noted the occurrence in contact zones of 
diploid and tetraploid toads (Borkin et al., 1997, 
2000). Preliminarily, Lattes (1997) and Cervella et al. 

(1997) indicated that triploids from Kyrgyzstan (Kok- 
jar, near 39; Castellano et al., 1998) do not originate 
from hybridizations between diploid and tetraploids 
but seem to be closely related forms of the tetraploids. 
Presumably all-triploid populations of green toads 
from the Karakoram range and Western Himalayas 
(46, 47, 48a) have been discovered by Stock et al. 

(1998) and were described as a new subspecies (Stock 
et al., 1999). The new data from Pakistan (47a, 48b-e) 
suggest that triploids, which form at least at some 
localities (e.g., 47) all-triploid gonochoristic popula- 
tions, are distributed along the upper tributaries of the 
Gilgit river (48b), they live at the Shandur pass (48c, 
3720 m a.s.l), and also in the Central Hindukush, i.e. 
the Chitral valley (48d, e). Interestingly, in the Kara- 
koram exists obviously an eastern limit of the distri- 
bution of triploids situated between their most eastern 
record (48a) and the most western one (49) of diploid 
B. latastii in the gorge of the Indus river. Based only 
on morphological characters, Baig (1998) reported on 
the sympatric occurrence of "Bufo latastii" and "B. 
pseudoraddei" in the "Neelam valley" (= Jhelum val- 
ley) of Azad Kashmir in eastern Pakistan. 

Generally, the correlation of distributional data on 
green toads with global climatic factors appears to be 
problematic because the local or microclimatic condi- 
tions to which the animals are adapted may differ 
strongly from the climatic zone in which any locality 
is situated. 

Zoogeographic implications. Borkin (1999: 350) 
considered "the B. viridis group as an indicator of 
southern limits of the Palearctic in arid regions". 
Mazik et. al. (1976) as well as Pisanets (1978) already 
reported an allopatric occurrence of diploid and tetra- 
ploid green toads because they have different ecologi- 
cal preferences. This hypothesis is principally 
supported by the present map but it seems more 
appropriate to call the ranges parapatric (Borkin et al., 
1997). Many authors (Pisanets and Shcherbak, 1979; 
Toktosunov, 1984; Borkin et al., 1986b, c; Borkin and 
Kuzmin, 1988) arranged the tetraploids of the arid 
foothill regions in contrast to those of the high moun- 
tains. In our opinion, it appears possible that the limit- 
ing climatic factors in the high mountains as well as in 



the more continental high plains are similar and are 
obviously suitable only for polyploid green toads. 

At the moment, the causes for the specific distri- 
bution pattern remain speculation. Kuzmin (1995: 
189, 1999) refers to the high degree of temperature 
tolerance of tetraploid green toads (-30°C to 45°C). In 
the high mountain habitats as well as in the continen- 
tal deserts strong daily and annual variations of tem- 
peratures as well as a high mean annual solar radiance 
(e.g. in the northern Asian deserts 2700-2800 h, in the 
interior Tien Shan > 2600 h, see Zlotin, 1997) might 
have caused a genetically caused selective advantage 
of the polyploid forms. This may also account for the 
missing of diploid toads in the east of the range where 
only tetraploids have been detected. The eastern 
boundary of the range of tetraploid green toads and 
the obviously parapatric occurrence of B. raddei in 
Northern China and Western Mongolia was discussed 
by Peters (1971: "B. viridis" for tetraploids), Borkin 
and Kuzmin (1988), and Stock (1998b). The most 
western known records of tetraploids (Kopet Dagh, 
Khorasan) seem to correspond with the region of tran- 
sition from winter-mild, summer-dry steppe climates of 
Middle Asia and Afghanistan to the winter-humid, 
summer-dry climates of Mediterranean type in Iran 
(Walter and Lieth, 1967; Miiller, 1996). The deserts of 
Central Iran with fewer than 100 mm rainfall/year and 
large sand or salt areas seem to separate the polyploids 
in the East of Central Iran (and Afghanistan?) from the 
diploids in the West. Although B. surdus may provide 
unexpected karyological data, at the moment, in spite 
of only few studies, records of polyploid green toads in 
the West of Iran appear to some extent improbable. 
This may rather result from the history of distribution 
than from the possible relationship between polyploidy 
and environmental selective pressure. 

The present image of the distribution of diploid and 
tetraploid toads possibly also corresponds to the experi- 
ence that (especially allopolyploid organisms may 
have advantages in regions with extreme or changing 
environments (Futuyama 1990: 69 - "polyploids are 
more resistant"), when colonizing new habitats 
(Bretagnolle et al. 1998) or were considered to have a 
higher competitive ability (e.g., Lumaret et al., 1997). 
However, particularly the distribution of diploid plant 
species and their polyploid relatives probably more 
strongly reflects the influence of historical factors than 
of their current ecological requirements (Bretagnolle et 
al. 1998) or adaptedness. Morescalchi (1990) stated: 
"In amphibians, changes in genome size (...), far from 
being random, they are related to metabolic and onto- 
genetic factors which are of crucial importance in the 
adaptive strategies of these animals". 



2001 



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Vol. 9, p. 91 



Acknowledgments 

We thank Prof. Dr. J. Martens, Mainz, for mating call 
recordings from northern Iran, and Dr. T. 
Klapperstiick, Halle, for using the CYDOK image 
analyses system. Prof. Dr. N. Ananyeva, St. Peters- 
burg, Dr. R. Giinther, Berlin, Dr. C. McCarthy, Lon- 
don, Dr. G. Nilson, Goteborg, Dr. J. B. Rasmussen, 
K0benhavn, for loaning preserved specimens. Chro- 
mosome photographs were kindly prepared by G. 
Hesse, Wurzburg. The basic computer-map was cre- 
ated by H. Nagel, Halle, according to our concept. 
Many thanks to Dr. K. J. Baig, Islamabad, Dr. L. Ya. 
Borkin, St. Petersburg, Dr. S. Castellano, Torino, Prof. 
Dr. A. Dubois, Dr. A. Ohler, Paris, Dr. T Papenfuss, 
Berkeley, J. F. Schmidtler, Munich, and Prof. Dr. E. 
Zhao, Chengdu, for help with getting literature. Dr. S. 
L. Kuzmin, Moscow, kindly allowed to use his data- 
base "Amphibians of the former USSR". 

M. Stock is very grateful to the family of M. and 
M. Khatiri, Gorgan, for their kind hospitality, R. 
Dressel, Dresden, for optimism and assistance during 
the field work in Pakistan, Dr. D. Lamatsch, 
Wurzburg, for help with flow cytometry, and T. Diet- 
erich, Greifswald, for a toad from Tengiz Lake (Kaza- 
khstan). 

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plex) in Middle Asia. Amphibia-Reptilia. 19:29-42. 

Stock, M. 1998b. Tetraploid toads (Bufo viridis com- 
plex) from north-western China and preliminary taxo- 
nomic conclusions for Bufo nouettei Mocquard, 1910. 
Zeitschrift fur Feldherpetologie (Bochum) 5:139-166. 

Stock, M. and W.-R. Grosse. 1997a. Erythrocyte size 
and ploidy determination in green toads {Bufo viridis 
complex) from Middle Asia. Alytes (Paris) 15:72-90. 

Stock, M. and W.-R. Grosse. 1997b. New data from 
the Bufo viridis complex in Central Asia. In Rocek, Z. 
and Hart, S. (eds.) Herpetology '97, Abstracts of the 
Third World Congress of Herpetolgy 2-10 August. 

1997. Prague, Czech Republic. 

Stock, M., W.-R. Grosse, M. Schmid and C. Steinlein. 

1998. Ploidy level diversity in the Bufo viridis com- 
plex of Central Asia Zoology (Jena). Suppl. 1:76. 

Stock, M., M. Schmid, C. Steinlein and W.-R. Grosse. 

1999. Mosaicism in somatic triploid specimens of the 
Bufo viridis complex in the Karakoram with examina- 
tion of calls, morphology and taxonomic conclusions. 
Italian Journal of Zoology 66 (3): 215-232. 

Toimastov, S. S. 1989. Osobennosti morfologii zhab 
iz okrestnostei oz. Jashil'kul' na Pamire. Izvestiya 
Akademii Nauk Tadzhikskoi SSR, otdelenie biolog- 
icheskikh nauk 4 (1 17):43-48. (In Russian). 

Toktosunov, A. T. 1984. Ecological basis of altitude 
adaptation of the vertebrates of Tian-Shan. Nauka 
Publ. House. Leningrad. (In Russian). 

Toktosunov, A. T. and E. Yu. Mazik. 1977. Napravlen- 
nost' zhab v Tyan' -Shane, pp. 203-204. In Darevski, I. 
S. (ed.) Voprocy gerpetologii. Nauka. (In Russian). 
Ullerich, F.-H. 1966. Karyotyp und DNS-Gehalt von 
Bufo bufo, B. viridis, B. bufo x B. viridis und B. 
calamita (Amphibia, Anura). Chromosoma 18:316- 
342. 

Walter, H. and S.-W. Breckle. 1986. Okologie der 
Erde. Vol. 4. Stuttgart. Gustav Fischer. 

Walter, H. and H. Lieth. 1967. Klimadiagramm- 
Weltatlas. 3rd. ed., Gustav Fischer, Jena. 

Wu Min and Zhao Yajiang. 1987. A preliminary study 
of the karyotype of Bufo viridis Laurenti in Xinjiang. 
Zoological Research, Kunming 8 (4):339-342. (In 
Chinese with English summary). 

Zhao, E.-M. and K. Adler. 1993. Herpetology of 
China. Oxford, Ohio, USA, SSAR:l-522pp. 



Zlotin, R. I. 1997. Geography and organization of 
high mountain ecosystems in the former USSR. pp. 
133-159. In Wiegolaski, F E. (ed.) Ecosystems of the 
world, vol. 3: Polar and alpine tundra. Elsevier, 
Amsterdam. 



Vol. 9, p. 96 



Asiatic Herpetologieal Research 



2001 



Apppendix 1 



L \£' Ploidy Topographic description Source Method 


1 


2n 


Iran. Zagros Mountains. Gholaman. 30 km W Khorram „„..„, _,„„ 
Abad. 33=25' N. 48° 1 2' E preSent paper 


erythr. size, microdens. 


la 


2n 


Iran. SW-slope of Zagors mountains. Khuzestan 
Province. Choqa Zanbil. 32°31'N. 48°32' E. 560 m 
a.s.l. 


present paper 


low cytom. 


2 


2n 


iran. N-slope of Elburz mountains, valley 15 km S 
Gorgan, approx. 1 100 m a.s.l. 


Stock (1995) 


chrom count, of larvae and 
uveniles 


2a 


2n 


Iran. S-slope of Elburz mountains. N of Theran. near 
Polur. approx. 2350 m a.s.l. 


Martens unpublished, present paper 


calls 


2b 


- Iran, S-slope of Elburz mountains, N of Theran, valley 
NE of Fasham. aporox. 2540 m a.s.l. 


Martens unpublished, present paper 


calls 




2n 


Iran, S-slope of Elburz mountains. Lar valley NE 


Andren and Nilson (1979). present paper 


calls 


2c 


2n 


Iran. Tehran Province, N environs of Tehran, garden 
of Plant Pests and Diseases Research Institute 


Borkin et al. (2000) 


flow cytom. 


2d 


2n 


Iran. Tehran Province, approx. 70 km W of Karaj. 
Karpuz-Abad village 


Borkin et al. (2000) 


flow cytom. 


3 


2n 


Iran, NE. frontier zone near Turkmenistan, approx. 50 
km NE Gonbad-e-Kavus, 250 m a.s.l.. 37°38' N, 55°29' 
E 


Stock (1997a), Stock and Grosse (1997a), 
Stock (1998a) 


chrom. count, of larvae, adults, 
erythr. size 


4 


4n 


Turkmenistan Nebil-Dagskii Rayon N-slope of Stflck ( , Qg7a) StSck wd Grosse ( 1997a) , chr om. count, of larvae, adults, 
Bolshoi Balkhan. approx. 15 km S of Oglanly village, stock (1998a) erythr size, calls 
500 m a.s.l., 39°43' N, 54°29' E 


5 


2n 


Turkmenistan. Ashgabadskava oblast near Danata 


Pisanets (1978), Borkinetal. (1986a) chrom. count 




2n 


-"- 


Mezhzherin and Pisanets (1990), Pisanets 
(1992a) 


ext morph. 




4n, 3n 




Pisanets (1978) 


chrom. count 




4n 


Borkinetal (1986a) 


chrom. count. 




4n 


Mezhzhenn and Pisanets (19901. Mezhzhenn 
and Pisanets ( 1991 ). Mezhzherin and Pisanets 
(1995a. b) 


chrom. count and/or ext morph. 




4n 


Ataev (1987), Pisanets (1992a) 


ext morph. 




4n 


1 Pisanets (1992b) 


chrom count, ext morph. 




4n 


Turkmenistan, Ashgabadskaya oblast, stream 2-4 km 
SE of Danata village and warm spring approx. 4 km SE 
of Danata. 200 m a s 1 


Stock ( 1997a), Stock and Grosse (1997a), 
Stock (1998a) 


chrom. count, of larvae, adults, 
erythr. size, calls 


7 


4n 


"South-west Kopet-Dagh", not exactly localized 


Pisanets and Vasilenko (1995) 


method not described 


6 


4n 


Turkmenistan, Kuruchsudon-Reserve, Kopet-Dag (not 
exactly localized) 


Borkinetal (1986a) 


chrom. count, and/or flow cytom. 


7 


2n 


Turkmenistan, Kyzyl-Arvatskii Rayon. Kopet-Dag- 
Range, valley approx. 25 km SW of the station Bami. S 
of the pass. 750 m a.s.l., 38°37' N. 56°38' E 


Stock (1997a). Stock and Grosse ( 1 997a), 
Stock (1998a) 


chrom. count of larvae, erythr. size, 
microdens. 


8 


2n 


Turkmenistan. Aydere oasis, 1 Kara-Kala 


Borkinetal. (1986a) 


chrom count and/oi flow cytom 




4n 


Turkmenistan. Adere-oasis. E Kara-Kala 


Borkinetal. (1986a) 


chrom. count and/or flow cytom. 




4n i Turkmenistan, Kara- Kala Mezhzhenn and Pisanets (1995a. b) 


chrom. count and/or ext. morph. 


9 


2n 


Turkmenistan, lake Sarykamysh Borkinetal (1986a) 


chrom. count 


9a 


2n 


„ ., ._ — ~ o ut. Schneider and Egiasarvan (1995), 
Kazahkstan. Guryev Town. Chornaya Rechka Dujsebaveva et al. ( 1997) 


calls 


10 


2n 


Turkmenistan. Ashgabadskava oblast. Bacharden Pisanets ( 1992a) 


ext morph. 




2n 


.". Mezhzhenn and Pisanets ( 1991 1 method not mentioned 




2n 


Turkmenistan, Ashgabadskaya oblast, Bacharden Mezhzhenn and Pisanets (1995a. b) chrom. count.and/or ext. morph. 




2n 


Turkmenistan. Ashgabadskaya oblast. S Bacharden, 
approx. 10 km W Kelyata, 500 m a.s.l., 38°14' N, 
57°31'E 


Stock (1997a). Stock and Grosse (1997a), 
Stock (1998a) 


chrom. count, of larvae, erythr. size, 
microdens. 


11 


2n 


Turkmenistan. Ashgabadskaya oblast near Ashgabad 


Borkinetal. (1986a) 


chrom. count 




2n 


Mezhzherin and Pisanets ( 1 990), Mezhzherin 
and Pisanets (1991 1 


ext. morph. 




2n 


Turkmenistan, Ashgabad, village Kodzh Mezhzhenn and Pisanets (1991) 


method not mentioned 




2n 


Mezhzhenn and Pisanets (1995a. b) 


chrom count and/or ext morph. 




2n 


Turkmenistan, Ashgabadskaya oblast near Ashgabad ! Pisanets and Vasilenko (1995) 


method not described 




2n 


Pisanets (1991) 


chrom. count 




2n Turkmenistan, Ashgabadskava oblast, Ashgabad Stock (1995) 


chrom. count, of larvae 




4n Turkmenistan, Ashgabad Roth and Rab (1 9861 


chrom. count. 


12 


4n 


Turkmenistan, village Kuruchaudan Pisanets (1987) 


clutch 


12a 


2n 


Iran, 20 km WNW Kapkan. Khorasan, 37°22' N, 
58°32'E, 1670 m a.s.l. 


present paper 


erythr. size, microdens. 


13 


4n 


Iran. 10 km NE Baghestan. Khorasan. N of Kuh-e- 
Kalat 34°09' N. 58°25' E. 1900 m a.s.l. 


Stock et al. (1998a). present paper 


chrom. count, erythr. size, 
microdens. and flow cytom 


13a 


4n 


Iran, Khorasan, Birjand, 32°33' N, 59° 10' N, about 
1500 m a.s.l. 


present paper 


chrom. count, flow cytom. 


13b 


2n 


Iran. Kerman Province, Kerman, 30° 18' N, 57°05' E, 
1860 m a.s.l. 


present paper 


chrom. count, flow cytom. 


13c 


2n 


Iran. Kerman Province, Bahr-e-Aseman Mountains, 
Sarduiyeh village in Sarduiyeh District 2500 m a.s.I. 


Borkin et al. (2000) 


flow cytom. 


14 


, Turkmenistan, lake Shach-Senem (not exactly 
zn [localized, SE of 9) 


Borkinetal. (1986a) 


chrom. count.and/or flow cytom 


15 


2n 


Turkmenistan, Ashgabadskaya oblast near Iolotan 


Mezhzherin and Pisanets (1990), Mezhzherin 
and Pisanets (1991) 


ext. morph. 


16 


4n 


Badchyz-Reserve, AkarChechme Pisanets (1978) 


chrom. count 




4n 


Borkin et al. (1986a) 


chrom. count 




4n 


_ " . 


Mezhzhenn and Pisanets (1995a, b) 


chrom. count.and/or ext. morph. 


17 


2n 


Uzbekistan. Buchara 


Roth and Rab (1986) 


chrom. count 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 97 



IS 


2n 


Uzbekistan. Samarkand Borkrn et al (1986a) chrom cuunl 


19 


4n 


Uzbekistan. Dzhisakskaya oblast Ravon Farislt Stock (1997a), Stock and Grossed 997a), chrom. count, of larvae, adults, 

Nurauu-Reserve N-slope r of Nuratau Range, 900 - | „ k , 9Qg ^ ■ ca , K 
1600 ma.s.l.. 40 35 N. 66 30 E | ' 




4n 


Uzbekistan, Dzhisakskaya oblast. Rayon Farish, S Stock (1997a), Stock and Grosse (1997a), chrom. count, of larvae, erythr. size, 
bank of Aidar-Kul near of Nuratau- Range, 300 m as. 1. Stock (1998a) microdens., calls 


20 


4n 


Tadzhikistan. 60 km E of mouth of Janob into 
Zeravshan river, right bank of Zeravshan river, near 
Pasucax 


Kryukovetal. (1985) 


chrom. count. 


21 


4n 


Tadzhikistan, Gissar- Range. Ziddi. 3000 m a.sl. 


Borkinetal. (1986a) 


chrom count 


22 


-In 


Tadzhikistan. Ramit-gorge, Chuhgaram 


Borkinetal. (1986a) 


chrom count 


4n Tadzhikistan. Ramit-gorge. Chuligaram Mezhzhenn and Pisanets ( 1995a, b) 


chrom. count and/or ext morph 


4n ■"- Pisanets (1992b) 


chrom. count., ext. morph 




, Tadzhikistan, 1 5 km SW of the mouth of Sardan- 
n Miena and Sorbo. gorge of Kafinugan, near Javroz 


Kryukovetal. (1985) 


chrom count. 


23 


2n and 4n 


Tadzhikistan, Dushanbe 


Roth and Rab (1986) 


chrom. count. 




4n Tadzhikistan, N Dushanbe 


Kryukovetal. (1985) 


chrom. count 


24 


4n 


Uzbekistan, Gissar- Range, 20 km of mouth of 
Tupalang-Darya. 800 m a si 


Borkinetal. (1986a) 


chrom. count. 


25 


2n 


Tadzhikistan, 90 km SW Dushanbe. Kafirnigan-valley 


Borkinetal. (1986a) 


chrom. count. 


26 


4n 


1 adzhikistan, 90 km S Dushanbe. Kafirnigan-valley. 
near lsambai 


Pisanets (1992b) 


chrom. count., ext. morph. 


27 


4n Tadzhikistan. S Pamir, near Lyangar 


Mezhzhenn and Pisanets (1990) 


ext. morph. 




4n Tadzhikistan. S Pamir, near Lyangar 


Mezhzhenn and Pisanets (1991) 


method not mentioned 




4n 1 -"-1800m a.sl Pisanets 1 1992b) 


chrom. count . ext. morph 


27a 


4n Tadzhikistan, near Faizabad Mezhzhenn and Pisanets (1991) 


method not mentioned 




4n I ad/hikistan. near Faizabad Pisanets (1991) 


chrom count 




4n 


Tadzhikistan. near Faizabad 


Mezhzhenn and Pisanets ( 1995a. b) 


chrom count. and/or ext. morph. 




4n 


- " - .localization using the information:„I20 km N of 
Dushanbe" in: 


Pisanets (1992a) 


ext. morph. 




4n 


. " _ 


Pisanets 1 1992b) 


chrom. count., ext. morph. 




4n 


ladzhikistan. Vakh valley, environs of Obi-Garm Mezhzhenn and Pisanets 1 1991 1 


method not mentioned 




4n 


Tadzhikistan. Obi-Gann. 2300 m a.s.l. Pisanets (1992a) 


ext. morph. 




4n 


ladzhikistan. Obi-Garm Mezhzhenn and Pisanets (1995a, b) 


chrom. count. and/or ext. morph 




4n 


. ". 


Pisanets (1992b) 


chrom count . ext. morph 


27b 


4n 


fadzhikistan. mouth of Komarou-river. 2000 ma.s.l. 
(not exactly localized) 


Borkinetal. (1986a) 


chrom. count. 


28 


2n 


Tadzhikistan. 10-12 km SW of the mouth of Vakhsh 
and Javansu. near Kurgan-Tiube 


Kryukovetal. (1985) 


chrom count. 




2n 


Tadzhikistan. 1 0- 1 2 km W of the mouth of Vakhsh and 
Javansu. left of Vachsh, Kuibyshevskii Rayon 


Kryukovetal. (1985) 


chrom. count. 




2n 


Tadzhikistan, 50 km NE of mouth of Vakhsh into 
Pvandzh. near Dusti 


Kryukovetal. (1985) 


chrom. count. 


29 


2n Tadzhikistan. Chirik 


Borkinetal. (1986a) 


chrom. count. 




2n 


environs of Chaartuz. Chirik 


Pisanets (1991 i 


chrom. count. 




2n 


Tadzhikistan, Beshkenlskre peski, valley of Kafinugan 
nver. environs of Chaartuz 


Mezhzhenn and Pisanets ( 1991) 


method not mentioned 




2n 


-"- 


Pisanets (1992a) 


ext morph. 


30 


4n 


Tadzhikistan, S of Aktau-Range, 80 km S Dushanbe 


Kryukovetal. (1985) 


chrom. count 


31 


4n 


Uzbekistan, Tashkent 


Borkinetal. (1986a) 


chrom count and/or flow cy torn. 




4n 


. " . 


Kudryavcev et al. (1988) 


cytophotometry 




4n 


. " . 


Roth and Rab (1986) 


chrom. count 




4n 


Uzbekistan, Tashkent, 450 m as.l., 41°16' N. 69°13' E 


Stock [1997a), Stock and Grosse (1997a) 
Stock (1998a] 


chrom. count of larvae, adults, 
erythr. size, calls 


31a 


2n Kazakhstan, environs of Chimkent 


Mezhzherin and Pisanets (1995a, b) 


chrom count and/or ext. morph. 




4n Kazakhstan, environs of Chimkent Mezhzhenn and Pisanets ( 1991) 


method not mentioned 


31b 


4n 


Kazakhstan. Kyzyl-kum desert, 50 km SW of Bairkurn DuJ , ehayeva e , al. ( 1997), Castellanoet al. 
village, Chimkent region. 250 ma.s.l. 67°25 N, 42 02 ,„-Ug ' 


chrom count 


31c 


4n 


Kazakhstan, Chimkent region. Jagbagly village, 1 100 
mas.l..70 i 32 , N,42°25'E 


Dujsebayeva et al. (1997), Castellano et al. 
(1998) 


chrom. count. 


32 


4n 


Uzbekistan. 80 km E Tashkent, entry of Chatkal- 
Reserve. 5 km SE Burchmulla, 900 m a.s.l., 41°35' N, 
70°07' E 


Stock ( 1 997a), Stock and Grosse (1997a), 
Stock (1998a) 


chrom. count. and/or erythr. size, 
calls 


33 


4n Uzbekistan. Kuraminsky-Range, 3000 ma.s.l. 


Borkin et al. (1986a) 


chrom. count 


34 


4n 


Tadzhikistan. S Pamir, near Khorog 


Mezhzherin and Pisanets (1990) 


ext morph. 




4n 


Tadzhikistan. Gorno-Badakhshan. S Pamir, near 
Khorog 


Mezhzherin and Pisanets (1991) 


method not mentioned 




4n 


- " - Mezhzhenn and Pisanets (1995a. b) 


chrom count and/or ext morph 




4n 


-"- Pisanets (1992a) 


ext morph 


35 


4n 


ladzhikistan, S Pamir, near Ishkashim Mezhzhenn and Pisanets ( 1990i 


ext morph. 




, Tadzhikistan, Gorno-Badakhshan, S Pamir, near 

Ishkashim 


Mezhzherin and Pisanets (1991) 


method not mentioned 




4n Tadzhikistan. S Pamir, near Ishkashim Me/hzhenn and Pisanets ( 1995a, b) 


chrom. countanoVor ext. morpjt 
chrom count 


36 


2n | Kazakhstan, Dzhambul Roth and Rab ( 1 986) 


36a 


2n 


(outside the map ) Kazakhstan. Tengis Lake. 1 2 km W Dieterich leg., examined by Stock, 
ol Abaya village, about 50 40 N 69 4(1 1 unpublished 


flow cytometry (one specimen) 


37 


4n 


Kazakhstan, desert near Burubaital, S Balchash lake 
region 


Egembergdieva ( 1983) cited by Borkin et al. 
(1986a) 


unknown, original paper not 
available 




4n ! Kazakhstan, desert near Burubaital,SBalchash lake Borkinetal (1995) flow cytometry 

region 



Vol. 9, p. 98 



Asiatic Herpetological Research 



2001 



37a 


4n 


Kazakhstan. Karaoi village. 350 m as 1 , 74 47' N. Dujsebayeva et al ( 1997). Castellano et al. 
45°54'E (1998) 


:hrom. count., calls 


37b 


4n 


Kazakhstan. Zhidely Channel. Hi River Delta, 370 m Dujsebayeva et al. (1997), Castellano et al. 
a.s.i.,75°12'N.45°18'E (1998) 


:hrom. count., calls 


38 


2d 


_,. „ ,„„ , Mazik. Kadyrova and Toktosunov (1976) and 
Kyrgyzstan, Chu-valley. 650 m a.s.l. C|ted bv Bo rkin et al . ( 1 9Sh., 1 


:hrom. count. 




2n 


Kyrgyzstan, Chu-valley Toktosunov and Mazik (1977) method not mentioned 




2n 


Kvfvzstan Tulek Castellano et al (1998) chrom count, calls 




2n 


Kyrgyzstan, Tyulyok, not exactly localized | Borkin et al. (1986a) chrom. count.and/or flow cytom. 


39 


2n 


Kyrgyzstan, Frunze (Bishkek) Bachmann et al. (1978) cytophotometry 




2n 


Borkin et al 1 1986a) 


chrom count, and/or flow cytom. 




3n 
4n 


Kyrgyzstan. S Bishkek Kuzmin(1995 187) 
Kyrgyzstan. Frunze (Bishkek) Borkin et al. (1986a) 


chrom. count. 




4n 


Kvrgvzstan. environs of Frunze (Bishkek) Mezhzhenn and Pisanets ( 1991 ) jmefhod not mentioned 




4n 


Bachmann et al. (1978) cytophotometrv 




2n 


Kyrgyzstan. Bishkek (Frunze) 


Stock ( 1 997a ), Stock and Grosse ( 1 997a) 


chrom. count.and/or erythr. size 


? 


4n 


Kyrgyzstan. Kyrgyzskn Khrebet, 1200 m a.s.l. and 
2500 m a.s.l.. not exactly localized 


Fikhtman(1989) 


method not mentioned 


9 


2n, 3n, 4n 


Kyrgyzstan. Kok-jar, 25 km S of Bishkek, approx. 
1300 m a.s.l., artificial bassin about 6 km from the 
villare 


Castellano etal. (1998) 


chrom. count., calls 


40 


4n 


Kyrgyzstan, Toktogul-valley. 900 m a.s.l. 


Toktosunov (1984) and cited by Borkin et al. 

(1986a) 


chrom. count. 


41 


2n? 


Kyrgyzstan, SW slope of Fergana- Range, near 
Arslanbob, 1 800 m a.s.1. 


Pisanets and Shcherbak ( 1979) 


method not described 




3n 


.". 


Stock (unpublished) 


chrom. count, (one juvenile animal) 




4n 


_ " . 


Borkin et al. (2000) 


method nut mentioned 


42 


4n 


Kyrgyzstan, Arpa-valley. 3500 m a.s.l. 


Toktosunov (1984) and cited by Borkin et al. 
(1986a) 


chrom. count. 


43 


4n 


China, Kashgar, 39°29'N, 76°02'E. 1350 m a.s.l. 


Stock (1998b) 


chrom. counl . flow cytom . and/or 
ervlhr size, calls 


44 


4n 


China, E-Pamir. Taxkurgan, 37°47'N, 75° WE, 3350 m 
a.s.l. 


Stock et al. (1998a), Stock (1998b) 


chrom. count., calls 


45 


4n 


Tadzhikistan, Central Pamir, lake Jashilkul. 3734 m 


Toimastov (1989) 


ext. morph. 




4n 


- " - (also 3n, 2n ?) Mezhzhenn and Pisanets ( 1 990) 


ext. morph. 




4n 


_ " - Mezhzhenn and Pisanets (1991) 


method not mentioned 




4n 


- " - (also 3n, 2n ?) Pisanets (1992a) 


est morph 




4n 


- " - (also 3n, 2n ?) Pisanets (1992b) chrom. count., ext. morph. 




4n 


- " - (also 3n, 2n ?) 


Mezhzhenn and Pisanets ( 1 995a, b) chrom. count.and/or ext morph. 


46 


3n 


Pakistan. Karakoram, Sust, Hunza-Valley, 36°46iN, 
74°50'E, 2950 m a.s.l. 


Stock etal. (1998. 1999) 


chrom. count., flow cytom., and/or 
erythr. size, calls 


47 


, Pakistan. Karakoram. Pasu, Hunza- Valley. 36°30'N, 
ia 74°52'E. 2600-2800 m a.s.l. 


Stock et al. ( 1 998, 1 999 ). present paper 


chrom. count., flow cytom., and/or 
erythr. size, calls 


47a 


3n 


Pakistan. Northern Areas, Hunza-Valley, river bank. 
Karimabad near Ganesh, 36°I8'N, 74°4FE, 2060 m 
a.s.l. 


present paper 


flow cytom. 


48 


3n 


Pakistan, W-Himalayas, Upper Swat valley. Kulalai, 
35° 1 9'N, 72°36'E, 1 750 ma.s.1. 


Stock etal. (1998, 1999) 


chrom. count, only one specimen), 
and/or erythr. size, calls 


48a 


3n 


Pakistan, Northern Areas, Gilgit, 35°54'N, 74°24'E, 
1550 m a.s.l. 


Stock etal. (1998, 1999) 


chrom. count., flow cytom.. and/or 
erythr. size, calls 


48b 


3n 


Pakistan, Northern Areas, Gupis, 36°14'N, 73°27'E, 
opposite Yasin valley, 2160 m a.s.l. 


present paper 


flow cytom. 


48c 


3n 


Pakistan, Shandur Pass, Lake, border of NWFrontier 
Prov. and Northern Arears, 3720 m a.s.l. 


present paper 


flow cytom. 


48d 


3n 


Pakistan, NWFrontier Prov.. Hindu-Kush, Buni, 
approx. 36°20'N, 72°20'E, approx. 1900 m a.s.l. 


present paper 


flow cytom. 


48e 


3n 


Pakistan, NWFrontier Prov., Hindu-Kush, Chitral City, 
35°53'N. 71°47'E, 1480 m a.s.1. 


present paper 


flow cytom. 


48f 


3n? 


Afghanistan. Kabul ?. 34°31iN, 69°12'E, sample not 
surely localized for Kabul 


Bachmann et al. (1978): "36% more DNA 
than diploid B. viridis ", Hemmer et al. (1978) 


microdens. 


49 


2n 


Pakistan, Northern Areas (Baltistan). Himalaya, 
Satpara river SW of Skardu, 35"17 - N, 75°37'E. 2300 
m a.s.1., see also M and N 


present paper 


flow cytom., chrom. count 


50(?) 


4n 


China, Xinjang; Hotan; localization in the present map 
not sure 


Whu Mm and Zhao Yajiang (1987) 


chrom. count. 


51 


4n 


Kyrgyzstan. Naryn 


Stock (unpublished) 


chrom. count. 


52 


4n 


Kyrgyzstan. Kara-Kudzhur-valley 


Toktosunov (1984) and cited by Borkin et al. 
(1986a) 


chrom. count. 


52-57 


4n 


Kyrgyzstan. whole environs of Issyk-Kul. 1670 m a.s.l 


Mazik etal. (1976) 


chrom. count 




4n 


.-. 


Toktosunov (1984) and cited by Borkin et al. 

(1986a) 


chrom. count. 




4n 


Roth and Rab ( 1 987b) , chrom. count. 




4n 


Borkin ( 1 989) partly chrom count. 




4n 


.". 


Mezhzhenn and Pisanets I !99S a . b) 
Fikhtman(1989) 


chrom. count.and/or ext. morph. 




4n 


.". 


method not mentioned 


53 


4n 


Kyrgyzstan, Chu-valley. approx. 20 km W Rybache, 


Stock (1997a), Stock and Grosse (1997a) 


erythr. size 




4n Kyrgyzstan. SW bank of Issyk-kul 


Me/hzhenn and Pisanets ( 1991 ) 


method not mentioned 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 99 



,, . Kyrgyzstan. Issyk-kul. N-bank near Sary-Kamysh. Stock ( 1997a), Stock and Grosse (1997a). —al- .:„ —n. 
34 n 1670 m a.s.l.. 42°29'N,76°20'E ISlock (1998a) eryuir. size, cans 




4n 


Kvrgvzstan. Issyk-kul [Castellano et al (1998) 


chrom. count., calls 


55 


4n 


Kyrgyzstan, Issyk-kul. S-bank nearTamga, 1670 m 
a.s.l 


Stock (1997a). Stock and Grosse (1997a). 
Stock (1998a) 


chrom. count, of adults, erythr. size, 
calls 


56 


4n 


Kyrgyzstan. Rayon Tyub. village I-runze 


Borkin (1989) 


partly chrom count 


57 


4n 


Kyrgyzstan. NO-bank of Issyk-Kul. Rayon Tyub. near 
Kuturga 


Borkin (1989) 


partly chrom. count. 




4n 


Kyrgyzstan. Issyk-kul. N-bank near Chon-Oryuktu. 
1670 m a.s.l. 


Stock (1997a). Stock and Grosse (1997a) 


chrom. count of adults, erythr. size 


58 


4n 


Kyrgyzstan. Kemin-valley. 2500 m a.s.l. 


Toktosunov (1984) and cited by Borkin et al. 
(1986a) 


chrom count. 


59 


2n 


Kazakhstan. Almaty 


Birstein (1981). Pisanets ( 1 99 1 ) 


chrom. count 




2n 


Kazakhstan. Almaty ] Mezhzhenn and Pisanets ( 1 99 1 ) 


method not mentioned 




4n 


Kazakhstan, Almaty Mezhzherin and Pisanets ( 1995a. b) 


chrom. count. and/or ext morph. 




4n 


Kazakhstan. Almaty Borkin et al. (1995) 


flow cytometry 




4n 


Kazakhstan. Almaty. 900 m a.s.l., 76°55' N, 43° 15' E 


Dujsebayeva et al. (1997). Castellano et al. 
(1998) 


chrom. count. , calls 


59a 


2n 


Kazakhstan, Kopa, 20 km S of Kopa station. 75"47' N. 
4< 25' E 


Dujsebayeva et al. (1997), Castellano et al 
(1998) 


chrom. count, calls 


59b 


4n 


Big Almaty Lake. Zailiskii Alatau Range. 2300 m 
as 1 , 77°N. 43°04' E 


Dujsebayeva et al (1997), Castellano et al 
(1998) 


chrom. count., calls 


60 


4n Kazakhstan. Kapchagav. Ili-river 


Borkin etal. (1986a). Borkin et al (1995) 


chrom. count and/or flow cytom. 


60a 


. Kazakhstan. Bashn, 1 km S Altyn-Emel Mountain 
Range 44' 10' N. 78 "45' E 


Borkin etal. (1995) 


flow cytometry 


60b 


4n S foothills of the Koyandytau Mountain Range 


Borkin etal. (1995) 


How cytometry 




, Ayan-Saz Point. Borokhudzir river valley, between 
Kovandvtau and Dsunganan Alatau Mountains 


Borkin etal. (1995) 


flow cytometry 


61 


-, ^ 4n Kazakhstan, valley of river Chingzhal, basin of lake 
' ' Alakol. 6- 1 km near Andreevka 


Golubev(1990) 


method not mentioned 




4n Kazakhstan. Andreevka 


Mezhzhenn and Pisanets (1995a. b) 


chrom count and/or ext. morph. 




4m Kazakhstan. Taldy-Kurganskaya oblast 


Pisanets 1 1992b) 


chrom. count., ext. morph. 




4n Kazakhstan. Ucharal Mezhzhenn and Pisanets (1995a, b) 


chrom. count and/or ext. morph. 


62 


4n China. Xinjang; Wusu (Usu ?) 


Whu Mm and Zhao Yajiang 1 1987) 


chrom. count. 


63 


4n 


China. E-Tian Shan (E-Narat Shan), Kunas 43°14'N, 
84 40'E, :i45 M , 


Stock (1998b) 


chrom. count., calls 


65 


4n 


Kazakhstan. Aksnr-Farm. Zaysan-lake 


Borkin etal. (1986a) 


chrom count 




4n 


SW-part of Zaysan-gorge, 1 2 km NW of settlement 
Maikapchagai near Aksiir-Farm 


Shcherbak and Golubev ( 1981) 


method not mentioned 


64 


4n 


China, Xinjang, Wulumuqui (Urumqi) 


Whu Min and Zhao Yajiang (1987) 


chrom. count 


66 


4n 


Mongolia, river Ded-Nariin-Gol, S-slope of Mongolian 
Altai. 2000 m a.s.l., 1 2 of the detailed map in: 


Borkin and Kuzmin (1988) 


mostly chrom count. 


67 


4n 


Mongolia, oasis Chug-Bulag, Bulgan-Somon, Chovd- 
Aimak 


Borkin etal. (1986b) 


chrom. count. 




4n 


corresponds to 1. 2 and 3 of the detailed map in 


Borkin and Kuzmin (1988) 


chrom. count. 


68 


4n 


Mongolia, Ml kin W of settlement Bulgan, ( :hovd 
Aimak. nver Bulgan-Gol 


Meyer (1991) 


ext. morph. 


69 


4n 


Mongolia, spring Chujten-Bulak. S-slope of Mongolian 
Altai, 2000 m a.s.l., spring Bayan-Mod, 15 km N of 
spring Icher-Tol, 1600 m spring Icher-Tol, 83 km NW 
of settlement Bulgan, 1 600 m a.s.l., spring Chavchig- 
Us or Chavchig-Bulag-spring Shara-Bulag 


Borkin and Kuzmin (1988) 


partly chrom. count. 


70 


4n 


Mongolia, river Uench-Gol, 46°n.B., 92°w. L., 1350 m 
a.s.l. ( 1 1 of the detailed map in: 


Borkin and Kuzmin (1988) 


partly chrom. count. 




4n 


Mongolia, spring Jaman-Usny-Bulag or Jaman-Us. 30 
km E of settlement Uench, 1 800 m a.s.l., 1 2 of the 
detailed map in: 


Borkin and Kuzmin (1988) 


partly chrom. count. 


71 


4n 


Mongolia, oasis Ushigiin-Us, Dsungarian Gobi, 90 km 
SW of settlement Bulgan. 9 of the detailed map in: 


Borkin and Kuzmin ( 1988), Orlova and 
Uteshev(l986) 


partly chrom. count 


72 


4n' 


Mongolia, spring Domdzhigiin-Us. Dsunganan Gobi. 
85 km S of settlement Bulgan. 10 of the detailed map 
in: 


Borkin and Kuzmin ( 1 988), Orlova and 
Uteshev(1986) 


partly chrom. count 


73 


4n 


China, Xinjang; Hami prefecture, 1.' km M 
Koumenzi. 2090 m a.s.l. 


Zhao and Adler( 1993) 


ext morph. 


74 


4n 


China. Xinjang. Hami Whu Min and Zhao Yajiang (1987) 


chrom. count 


A 


? 


Iran, l.unstan. Shah Bazan. km 324 of the Transiranian ^h™*' ' n o^m'' ^^q,,^- „ 
•, .„ , , , e D , , , ,. ■ Schmidt er (1969 , Mertens 1971b), Eiselt 
railway, type locality of Bufo {surdus ) lunsmmcus a[)d Schmidtler ( , 973 . see als0 ^ s 


no ploidy determination 


B 


2n? 


Iran. NW pan of Central Iranian Plateau. Cheshmeh-ye 
Sefied-Ab, type locality of Bufo ku\ir?n.sis 


AndnSn and Nilson (1979), Stock (1998a): 
diploid'?, present pajper 


calls 


C 


2n 


Iran, Kerman, type locality of Bufo viridis 
kermunensi* 


Eiselt and Schmidtler ( 1 97 1 ), Hemmer et al. 
(1978), present paper 


chrom. count, flow cytometry 


D 


4n 


Turkmenistan, Achgabadskaya oblast, Danata village, 
type locality of Bufo danaiensis 


Pisanets (1978). Borkin and Kuzmin (1988), 
see also number 5 


chrom count 


E 


4n 


E-Iran, Ssaman Shakhi mountains near Birjand, type 
locality of Bufo oblongus 


Nikolsky (1896 1897), Eiselt and Schmidtler 
(1973). Roth (1986). Borkin and Kuzmin 
( 1 988). present paper 


chrom. count, flow cytometry 


F 


2n 


Hemmeret al. (1978). Pisanets and Shcherbak 
Tadzhikistan. Dushanbe, type locality of Bufo i in dis (1979), Roth (1986), Borkin etal. (1986b), no ploidy determination of the type 
turanensis Kuzmin (1995, 1999), Mezhzherin and series 

Pisanets (1995a. b). see also number 23 



Vol. 9, p. 100 



Asiatic Herpetological Research 



2001 



c -, Tadzhikistan, SW-part Beshkent desen, near Shaartus. PisanetM 1991 1, Pisanets el al (1996), •aasA 
type localitv of Bufo shaarmsiensis Kuzmin (1999). see also number 29 


H 


? 


„ .. . ~ .. , , ... rn_r / • -j- \ Eiselt and Schmidller ( 1 973 ). Borkin and 
Pakistan, Pishin, type locality of Bufo (vmdts ) Kuzmn , 98g S(6ck ■ a , 999 Borkl[) e( 

zugmayen 'al.(2000) 


no ploidy determination 


J 


3n and/or 
4n(?) 


Pisanets and Shcherbak (1979: diploid 
Kyrgyzstan. Arslanbob. type locality of Bufo viridis without determination), Borkin and Kuzmin 
asiomontanus (1988), Borkin et al. (1997), Kuzmin (1999), 

Borkin et al. (2000) 


chrom. count, (only one triploid 
specimen tested by Stock, unpubl.l. 
calls 


K 


4n 


„ „ _. . . , „ . ,„, Kashchenko(1909). Stock (1997a), Stock and 
Kyrgyzstan, Kokmoinok. terra typica of Bufo Qws%e , 99? R mjn , 999 see ^ chrQm 

(vind«)iim<:otor number 52 to 57 


L 


4n 


China. Kashgar. one of three type localities of Bnjo Mocquard (1910). Borkin and Kuzmin (1988), 
nouenei sensu Mocquard; type locality of the B Stock ( 1 998b), additional papers are cited in 
nouettei lectotvDe the last one 


chrom. count., calls, flow cytometry 


M 


2n 


Pakistan, Shinu village near Siachen glacier, type 
locality of Bufo siachinensis 


Khan ( 1 997 ). Baig ( 1 998 ), Stock et al. ( 1 998, 
1999): junior synonym of B. latastii. see N 
and 49 


chrom. count., flow cytometry 


N 


2n 


N-India. Ladakh. a locality was not exactly described, 
terra typica of Bufo latastii 


Boulenger (1882: 295), Dubois and Martens 
(1977), Hemmer et al. (1978), Pisanets and 
Shcherbak (1979), Gruber (1981 ), Borkin et 
al. (1986a). Roth (1986). Borkin and Kuzmin 
(1988), Khan (1997). Baig (1998), Kuzmin 
( 1999), Stock et al. (1999). see M and 49 


chrom. count, flow cytometry 


O 


3n 


Pakistan, Swat valley, Mingora province, terra typica 
of Bufo (viridis ) pseudoraddei pseudoraddei 


Mertens (1971a), Pisanets and Shcherbak 
(1979). Roth (1986), Borkin and Kuzmin 
(1988), Baig (1998), Kuzmin (1999), Stock et 
al. (1999), additional papers are cited in the 
last one 


chrom. count, (only one specimen). 
calls 


P 


3n 


N-Pakistan, Karakoram, type locality of Bufo 
pseudoraddei baturae 


Stock etal. (1999) 


chrom. count., calls, flow cytometry 


Q 


? 


SE-Iran, Bazman, Tamin in Sarhad, Duz-Ab. Ziaret in 
Sarhad. type locality of Bufo persicus 


Nikolsky ( 1 899). Carvsky ("1925". 1 926): 
"synonym of B surdus ", Schmidt (1955), 
Schmidtler and Schmidtler ( 1969). Eiselt and 
Schmidtler (1973), see also R and S 


no ploidy determination 


R 


? 


Belutchistan (W-Pakistan?), not exactly localized, terra f^X'a'n^Schrnidt'ler ( 1969?. Eisllt^d 
typica of Bufo surdus Schmidtler ( 1 973). see also Q and S 


no ploidy determination 


S 


7 


SW-lran. Mehkuh. 70 km S of Shiraz, type locality of Schmidtler and Schmidtler (1969), Eiselt and 
Bufo surdus annulatus Schmidtler ( 1 973) 


no ploidy determination 


T 


4n 


China, type locality of Bufo ssp. taxtorensis 


Fei et al. (1999), Stock (1998b) 


no ploidy determination in ssp. 
description, Stock (1998b): chrom 
count, (only one specimen), calls 


U 


7 


China, Kok-Far (= Kokyar). type locality of the 
depicted specimen of Bufo viridis var. pewzowi , three 
other type localities for the series 


Bedriaga (1898: 61 and Fig. 2. plate I), 
Hemmer et al ( 1 978). Borkin and Kuzmin 
(1988), Fei etal. (1999) 


no ploidy determination 


V 


7 


China. Tschik-Tym (= Qiktim). Turfan, type locality of 
Bufo viridis var. grum-grzinuiiloi ; "Turfan": one of 
14 type localities of Bufo viridis var. strauchi 


Bedriaga (1898: 61). Hemmer etal. (1978). 
Borkin and Kuzmin ( 1 988 ) 


no ploidy determination 



2001 



Asiatic Herpetological Research 



Vol.9, pp. 101— 106| 



First Record of the Smooth-Backed Parachute Gecko Ptychozoon lionotum 
Annandale 1905 from the Indian Mainland 

Samraat Pa war' and Sayantan Biswas 2 

Wildlife Institute of India, Chandrabani, Dehradun- 248 001, India. Present addresses: '3/21, Mohanwadi, 
AlandiRoad, Yerawada, Pune-4 11006, India. 2 30/3 Jheel Road, Calcutta-70003 1 , India. 

Abstract.- The smooth-backed parachute gecko, Ptychozoon lionotum is reported from the mainland India for the 
first time. The nearest known previous record was from Pegu, Myanmar, about 700 km southeast of the previous 
location. The species was collected in Langtlai and seen in the Ngengpui Wildlife Sanctuary, both in south 
Mizoram. The collected individual was kept in captivity for four and a half months, during this time, 
opportunistic observations on activity pattern, food habits, escape and parachuting behavior were made. Both 
individuals showed slow, deliberate pre-escape movement previously unrecorded for Ptychozoon. Information 
on morphological characters and morphometric measurements is presented. Explanations for the disjunct 
distribution are discussed. 



Key words.- Reptilia, Gekkonidae, Ptychozoon, parachute gecko, Northeast India, Myanmar, distribution, 
biogeography, behavior 



Ptychozoon is a genus of arboreal geckos distributed 
over much of Southeast Asia, primarily in moist tropi- 
cal evergreen and semi-evergreen forests (Brown, 
1999; Brown et al. 1997; Smith, 1935). At present, six 
species are recognized under the genus: Ptychozoon 
kuhli, P. horsfieldii, P. lionotum, P. intermedium, P. 
rhacophorus and P. trinotaterra (Brown et al., 1997; 
Brown, 1999). To date, the only species reported for 
India is Ptychozoon kuhli, from the Nicobar Islands 
(De Rooij, 1915; Smith, 1935; Tiwari, 1961). We 
present here the first record of the smooth-backed 
parachute gecko Ptychozoon lionotum based on two 
records from the state of Mizoram (21°56'N to 
24°31'N and 92°16'E to 93°26'E) in Northeastern 
India (Fig. 1). 

The first individual was sighted on 29th June 1998 
in Lawngtlai town of south Mizoram during a short 
survey. Subsequently, on 21st April 1999, a second 
individual was sighted by SP from the vicinity of 
Ngengpui Wildlife Sanctuary (NWLS; 22°21'24" N 
to 22°30'06" N and 92°45'12" E to 92 o 50'20•' E) in 
south Mizoram, during a herpetofaunal community 
study (Pawar, 1999). We could only collect the first 
individual, and although SP could get a superficial 
look at the second one before it escaped (see below), 
we presume that it was the same species as the 
straight-line distance between the two sites is only 
about 40 km. Both the localities lie in the low to mid- 
elevation region of south Mizoram. The vegetation is 
of the tropical (moist) evergreen type, corresponding 
to Northern Tropical Evergreen Forest (lb/c2; Cham- 



20° N 




18°N 

BAY OF BENGAL 



16° N 

50 100 150lrilom«OT 



96° E 



Figure 1. Present record (1) of Ptychozoon lionotum 
(BNHM 1445) from south Mizoram (India), along with 
nearest previous record (2) from Pegu (Myanmar). 



Vol. 9, p. 102 Asiatic Herpetological Research 2001 



Table 1 . Mensural (in mm) and meristic measurements of two specimens of Ptychozoon lionotum . The 
vouchered record reported here (BMNH 1445) is compared with one of the syntypes (ZSI 2601). The latter spec- 
imen was fully discolored and severely mutilated so many characters were not discernable (NA) and so measure- 
ments beyond mm were not possible. 



Character 

Head length 


BNHM 1445 
16.8 


ZSI 2601 

15 


Head width 


16.8 


16 


Head height 


12.7 


11 


Eye diameter 


4.8 


NA 


Eye to nostril distance 


8.2 


NA 


Eye to snout distance 


11.3 


11 


Eye to ear distance 


7.7 


8 


Inter orbital distance 


10.3 


NA 


Inter narial distance 


3.7 


NA 


Tympanum diameter 


2.3 


3 


Neck length 


11.9 


NA 


Snout to forelimb length 


36.0 


NA 


Axilla to groin length 


47.0 


42 


Body flap width (Greatest width from base of flap to tip) 


8.1 


9 


Body flap length (From axilla to groin) 


39.6 


41 


Fore arm length 


18.7 


12 


Fore limb length 


27.8 


21 


Femur length 


13.6 


NA 


Tibia length 


12.7 


NA 


Hind leg length 


22.9 


20 


Hind foot length 


39.6 


37 


Length of I st Toe 


6.3 


NA 


Length of IV th Toe 


9.4 


NA 


Snout to vent length 


94.6 


88 


Tail length 


93.0 


NA 


Tail width 


7.6 


NA 


Tail depth 


6.3 


NA 


Terminal tail flap length 


20.1 


NA 


Terminal tail flap width 


10.9 


NA 


Supralabials 


10/11 


NA 


Infralabials 


9/9 


NA 


Transverse dorsal bands in the axilla-groin region 


4 


NA 


Number of lobes fused before straight flap 


7 


NA 



2001 


Asiatic Herpetological Research 


Vol. 9, p. 103 




Character 

Supranasals in contact 




BNHM 1445 

No 


ZSI 2601 

NA 


No. of tail lappets 




19 


NA 


Subdigital lamellae (L/R) 


Finger 1 




11/10 


NA 


Finger II 




11/13 


NA 


Finger III 




12/16 


NA 


Finger IV 




15/15 


NA 


Finger V 




14/14 


NA 


Toel 




11/11 


NA 


Toe II 




12/12 


NA 


Toe 111 




16/15 


NA 


Toe IV 




14/14 


NA 


ToeV 




14/14 


NA 



pion and Seth. 1 968) and Chittagong Tropical Ever- 
green Forest ( lb/c4; Wikramanayake et al. 1998). 

Identification of the species as P. lionotum is based 
on the presence of the following combination of char- 
acters: absence of enlarged tubercle on the dorsum; 
denticulate tail lobes of the tail directed somewhat 
backwards; tail not tapering (Smith. 1935); presence 
of pre-digital notch on the forearm skin fold (Brown. 
1999; Brown et al. 1997; Cox et al. 1998). We also 
compared our specimen with one of the syntypes of P. 
lionotum (ZSI 2601. from Pegu. Myanmar) and a 
specimen of P. kuhli (ZSI 2603. from Nicobar Islands) 
housed at the National Zoological Collection of Cal- 
cutta. Selected morphometric measurements (after 
Brown, 1999; Brown et al. 1997; Das. 1997; Ota. 
1989; Zug and Moon. 1995) were recorded using 
Mitutoyo Digimatic callipers, with an accuracy of 0.1 
mm (Table 1 ). 

The color pattern of the specimen (in life) was as 
follows (Fig 2): Dorsally medium to dark gray with 
darker markings; dermal appendages lighter in color, 
mottled: distinct wavy dark, grayish-brown transverse 
bands present, four between the axilla and groin: chin 
and gular region dirty white to yellowish, white on 
chest, belly and underside of tail heavily powdered 
with gray-brown: underside of thighs, arms and der- 
mal appendages was similar to gular region in color. 
The collected individual showed some degree of color 
change in captivity, ranging from light (bands dis- 
tinct) to dark gray (bands barely distinct). 

The individual was judged to be an adult female 
based on the absence of preanal and femoral pores 



and the lack of hemipenal swellings at the tail base. 
The sex was later confirmed by dissection. 

The individual was kept in captivity in a glass ter- 
rarium measuring 2x1x1.5 ft, for four and a half 
months. During this period, we frequently took the 
gecko out at different times of the day, which allowed 
us to make additional behavioral observations, includ- 
ing those on its escape and parachuting behavior. The 
specimen was later euthanized and preserved in 70 9c 
ethanol after fixing in 10% formalin, and deposited in 
the reptile collection at the Bombay Natural History 
Museum (BMNH fide Leviton et al., 1985) in Mum- 
bai (No. 1445). 

Natural History and Behavior 

A mosaic of bamboo-dominated patches, remnant 
mature forest, teak plantations and jhum fallows of 
varying ages surround the town of Lawngtlai (900- 
1000 m elevation), where the first individual of Pty- 
chozoon was seen. At 1930 hrs on 29th June 1998. the 
parachute gecko was seen in a circuit house situated 
in the outskirts of the town. It was resting on the 
inside ledge of a window in the corridor, at a height of 
about 2 m from the floor. The corridor was enclosed, 
the only entry points being the windows and the doors 
al the ends of the passage. In the same passage, there 
were a few Hemidactylus frenatus, while a nearby 
corridor was occupied by Cosymbotus platyurus. 
While resting, the dermal appendages of the parachute 
gecko were closely apposed to the body, and it did not 
show any movement, except for a vigilant but slug- 
gish lateral movement of the body towards the outside 
of the ledge when attempts were made to capture it. 



Vol. 9, p. 104 



Asiatic Herpetological Research 



2001 




Figure 2. Ptychozoon lionotum (Adult female, BNHM 
1445) from Mizoram, northeast India. 

On 6th April 1999. SP, along with his field assis- 
tant, spotted the second Ptychozoon at 1820 hrs, next 
to a dirt track in a patch of mature evergreen forest 
south of NWLS houndary. -40 km (straight-line) 
south of the first locality. NWLS is the only remaining 
patch of unfragmented. mature primary forest in the 
area, characterized by a three-tiered structure, with 
towering, buttressed, deciduous emergents up to 50- 
60m in height, followed by middle and tertiary can- 
opy trees (Pawar. 1999). This area, especially the 
Ngengpui valley, experiences five rainless months, but 
the effective dry period is much shorter, with humid- 
ity being consistently high during these months due to 
fine, localized precipitation from cloud and fog. This 
individual was smaller than the first one and was spot- 
ted at a height of 5 m on the trunk of a Sterculia 
scaphigera tree. The tree is characterized by a deeply 
fluted trunk and a smooth but slightly flaking bark, 
and occurs as a deciduous canopy-emergent in pri- 



mary evergreen forest above 500 m elevation. The 
patch of forest was on a slope at an altitude of approx- 
imately 450 m. and the tree (385 cm in girth at shoul- 
der height) was towards the edge of the patch, slightly 
down slope, about 3 m from the dirt track and the 
observers. The gecko was sitting on the outer ridge of 
one of the trunk folds with its head pointing down- 
ward, barely visible on the lichen-covered bark. It was 
twilight, and upon sighting it. SP observed the animal 
for about a minute with the aid of a torch and binocu- 
lars before attempting to capture it. Meanwhile, the 
gecko had apparently become wary and steadily 
started moving laterally in the manner of the first indi- 
vidual away from the two observers, towards the other 
side of the trunk and out of sight. When SP tried to 
dislodge the gecko with a bamboo pole, the animal 
rapidly moved further around the trunk. It then 
lumped onto some lianas which were 2.5 m from the 
trunk, landed 1.5 m lower than its previous position 
with its head up. ran further up and vanished into a 
mass of dry branches which were caught in the lianas. 
All further attempts to trace the gecko were futile, and 
we presume that either the gecko jumped on to 
another tree or liana, or fell to the ground somewhere 
down slope when the lianas were shaken to dislodge 
it. Hemidactylus frenatus, H. garnoti, Cosymbotus 
platyurus and Gekko gecko are four other gekkonines 
that were commonly seen in the same area. 

During its four months of captivity, the individual 
was offered a variety of insects, of which it took cock- 
roaches and moths most readily. In the daytime, it 
remained motionless, either on one of the branches in 
its terrarium. or on one of the tar strips at the corners 
of the enclosure. Towards evening however, it would 
become active, and was often observed moving 
around the terrarium. making audible leaps across the 
corners of the enclosure. When taken out. its behavior 
was very different during day and night. If kept on a 
branch or tree trunk in the daytime, it would remain 
motionless with its limbs closely apposed to the trunk, 
and move only if provoked. If not disturbed for a long 
time, it would start moving slowly with the same 
slow, deliberate movement that it had displayed dur- 
ing its capture, either out of sight to the other side of 
the branch/trunk, or run up the tree. In the evenings 
however, it showed much more alacrity in trying to 
escape, often with the same preliminary lateral move- 
ment of its body. On two occasions it also resorted to 
launching itself into the air, and when it fell to the 
ground, remained motionless. This escape behavior 
has been earlier observed in these geckos, and remain- 
ing immobile ostensibly makes it difficult to locate 
them (Brown et ai, 1997). However, the slow pre- 



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Vol. 9, p. 105 



escape movement that we observed in both the speci- 
mens has not been reported before, and we reason that 
this behavior probably aids the gecko to position itself 
for parachuting or simply move out of sight (such as 
the blind side of a tree trunk) inconspicuously, with- 
out disclosing its crypsis. To gain further insights into 
this escape behavior that we observed in both the 
specimens, we dropped the individual on seven occa- 
sions from heights of 3-5 m and observed its behav- 
ior. In all instances, the dermal appendages came into 
play apparently due to air resistance, and the gecko 
dropped relatively softly on the ground without any 
horizontal displacement. 

The ecology and behavior of species of Pty- 
chozoon is poorly known, and there has been much 
discussion about its alleged ability to "glide" 
(Gunther, 1864; Smith, 1935; Tiwari, 1961 and refer- 
ences therein). It has been argued that the dermal 
appendages do not help in gliding, but enhance its 
camouflage. However, experimental studies have now 
demonstrated that the dermal appendages do allow the 
gecko to take advantage of air resistance while mak- 
ing long sallies (Heyer and Pongsapipatana, 1970; 
Marcellini and Keefer, 1976) and may serve a dual 
function in crypsis and escape or locomotion (Marcel- 
lini and Keefer, 1976). Recently, Brown et al. (1997), 
based on their observations of the escape behavior of 
P. intermedium in the wild, have argued that "para- 
chuting" is a more appropriate term than "gliding" to 
describe this behavior in these geckoes. Our observa- 
tions apparently sustain the arguments of Brown et al. 
(1997). Further studies on the preflight behavior of 
Ptychozoon species may provide interesting insights 
into the escape behavior of this extraordinary group. 

Biogeographical Notes 

The syntype (ZSI 2601 ), collected by Major Beddome 
and W. Theobald from Pegu in south Myanmar, was 
previously the northwestern most distribution record 
of Ptychozoon (Annandale, 1905; Brown et al. 1997). 
The present record thus adds a crucial link to the dis- 
tributional information for the genus, and increases 
the known range ca.700 km towards the northwest. 
This also adds another case of range disjunction in a 
region that already has numerous examples of taxa 
showing dramatic discontinuities in their range (Mani, 
1974). 

After the collision of the Indian plate with the 
Asian mainland in the Eocene (54-36 mybp) (Molnar 
and Tapponnier, 1975), Indo-Malayan faunal and flo- 
ral elements have colonized different parts of the 
India, resulting in more Indo-Malayan faunal repre- 
sentatives within India than vice versa (Das, 1996; 
Mani, 1974). Geckos are notorious for their penchant 



for waif-dispersal (Case et al. 1994). It has been 
observed that Ptychozoon species are not obligate for- 
est dwellers (Annandale, 1904; Brown et al. 1997), 
and probably are capable of dispersing through both 
forest and human inhabited areas (Annandale 1904; 
Brown et al. 1997). 

Mizoram is dominated by the Lushai Hills, a 
series of parallel hill ranges running from north to 
south and increasing in elevation from west to east 
(Pachuau, 1994). To the west of these hills lie the 
Chittagong Hill tracts of Bangladesh, and to the east 
lie the Chin Hills and the Arakan Yoma mountain 
ranges of Myanmar. The latter, also running in a 
north-south direction, lie between the lowland moist 
evergreen forests of south Mizoram and Pegu. Along 
the foothills of the Arakan Yomas, flanking the west- 
ern side, lie more or less contiguous rainforests, 
which forms a habitat bridge between these two areas 
(Collins et al. 1991). It is likely that this species has 
extended its range northwards along this route. This 
conjecture will get firmer footing if surveys in these 
forests reveal the presence of P. lionotum along these 
tracts. The areas beyond the Lushai Hills of Mizoram 
and the adjoining Chin Hills of Myanmar in contrast, 
are more arid with relatively dry forests. Moreover, 
recent surveys have not revealed the presence of Pty- 
chozoon species in these areas, and it is unlikely that it 
exists there (George Zug, pers. comm.). 

Our inquiries revealed that not many local people 
know of this gecko, but those who did, opined that it 
was rarely seen because it mostly "lived high up in the 
trees". There have also been unconfirmed reports of a 
parachute gecko from north Mizoram (Lai 
Ramthanga, pers. comm.). That this area has been 
inadequately surveyed is evident from the fact that the 
six-month herpetofaunal study conducted by SP 
yielded a number new taxa and distributional records 
(Pawar, 1999). Further exploration will probably 
reveal that Ptychozoon is present in other parts on this 
region, and its range may not be as disjunct as it 
appears now. 

Acknowledgments 

The Wildlife Preservation Society of India and Wild- 
life Institute of India supported our surveys in Mizo- 
ram. We are particularly grateful to the Mizoram 
Forest Department for permits and their support in the 
field. Zokhima was more than a field assistant to us. 
J.B. Alfred, S.K. Chanda and S.K. Talukdar at ZSI, 
Calcutta, kindly permitted us to access the collection 
and K. Deuti, I. Das and N.C. Gayen helped us to 
trace the specimens. R.M. Brown, I. Das, A. Sinha, 
N.M. Ishwar and K. Vasudevan provided us with criti- 



Vol. 9, p. 106 



Asiatic Herpetological Research 



2001 



cal comments and literature. Aysegul Birand helped 
prepare the map for Fig. 1 . 

Literature Cited 

Annandale, N. 1904. Contribution to Oriental Herpe- 
tology I: The Lizards of the Andamans, with the 
description of a gecko and a note on the reproduced 
tail in Ptychozoon homalocephalum. Journal of Asi- 
atic Society of Bengal 73:12-22. 
Annandale, N. 1905. Notes on some Oriental Geckoes 
in the Indian Museum, Calcutta, with the Description 
of New Forms. Annual Magazine of Natural History 
7:26-32. 

Brown, R. M. 1999. New species of parachute gecko 
(Squamata: Gekkonidae: Genus Ptychozoon) from 
northeastern Thailand and central Vietnam. Copeia 
1999(4): 990-1001. 

Brown, R. M., J. W. Ferner, and A. C. Diesmos. 1997. 
Definition of the Philippine Parachute gecko, Pty- 
chozoon intermedium Taylor 1915 (Reptilia: Squa- 
mata: Gekkonidae): redescription, designation of a 
neotype and comparisions with related species. Her- 
petologica 53:357-373. 

Case, T J., D. T. Bolger, and K. Petren. 1994. Inva- 
sions and competitive displacement among house gec- 
kos in the tropical Pacific. Ecology 75(2):464-477. 
Champion, H. G. and S. K. Seth. 1968. A revised sur- 
vey of the forest types of India. Government of India 
Printing, Delhi. 404 pp. 

Collins, N. M., J. A. Sayer, and T C. Whitmore (eds.). 
1991. The Conservation Atlas of Tropical Forests: 
Asia and the Pacific. IUCN, Macmillan Press Ltd., 
London. 256 pp. 

Cox, M. J., P. P. van Dijk, J. Nabhitabhata, and K. 
Thiraklupt .1998. A Photographic guide to snakes and 
other reptiles of Thailand and South Asia. Asia Books, 
Bangkok. 142 pp. 

Das, I. 1996. Biogeography of the reptiles of south 
Asia. Krieger Publishing Company, Florida. 87 pp. 
Das, I. 1997. Rediscovery of Lipinia macrotympamim 
(Stoliczka, 1873) from the Nicobar Islandsjndia. Asi- 
atic Herpetological Research 7:23-26. 
De Rooij, N. 1915. The reptiles of the IndoAustralian 
Archipelago, Vol. I and II. E.J.Brill, Leiden. 334 pp. 
Giinther, A. C. L. G. 1864. The Reptiles of British 
India. Ray Society, London. 452 pp. Reprinted ca. 
1982, Oxford and IBH Publishing Co., New Delhi. 
Heyer, W. R. and S. Pongsapipatana,. 1970. Gliding 
speeds of Ptychozoon lionotum (Reptilia: 



Gekkonidae) and Chrysopelea ornata (Reptilia: Colu- 
bridae). Herpetologica 26: 317-319. 

Leviton, A. E., Gibbs Jr., R. H., and Dawson, C. E. 
1985. Standards in herpetology and icthyology. part I. 
Standard symbolic codes for institutional resource 
collections in herpetology and icthyology. Copeia 
1985(4):802-821. 

Mani, M. S. 1974. (eds.) Ecology and Biogeography 
in India. Dr. W. Junk b. v. Publishers, The Hague. 773 
pp. 

Marcellini, D. L. and Keefer, T.E. 1976. Analysis of 
the gliding behavior of Ptychozoon lionotum (Rep- 
tilia: Gekkonidae). Herpetologica 32(3):362-366. 
Molnar, P. and P. Tapponnier. 1975. Cenozoic tecton- 
ics in Asia: Effects of a continental collision. Science 
1 89(4201 ):419-426. 

Ota, H. 1989. Japalura breviceps Gressit (Agami- 
dae:Reptilia), A valid species from a high altitude area 
of Taiwan. Herpetologica 45(l):56-60. 
Pachuau, R. 1994. Geography of Mizoram; R.T 
Enterprise, Aizawl. 153 pp. 

Pawar, S. 1999. Effect of habitat alteration on her- 
petofaunal assemblages of evergreen forest in Mizo- 
ram, North-East India. M.Sc. Thesis. Saurashtra 
University. 64 pp. 

Smith, M. A. 1935. The Fauna of British India, Cey- 
lon and Burma: Amphibia and Reptilia, Vol.11. - Sau- 
na, Vol. II. Taylor and Francis Ltd., London. 240 pp. 
Tiwari, K. K. 1961. The eggs and flight of the gecko 
Ptychozoon kuhli from Car Nicobar. Journal of Bom- 
bay Natural History Society 58(2):523-527. 
Wikramanayake, E. E. Dinerstein, P. Hedeo and D. 
Olson. 1998. Terrestrial Eco-regions of the Indo- 
Pacific Map, Publ. by WWFAVCMC, Washington 
D.C., USA 

Zug, G. R. and B. R. Moon. 1995. Systematics of the 
Pacific slender-toed geckos, Nactus pelagicus com- 
plex: Oceania, Vanuatu and Solomon populations. 
Herpetologica 5 l(l):77-90. 



2001 



Asiatic Herpetological Research 



Vol.9, pp. 107-112 



A New Species of Eremias (Sauna: Lacertidae) from Highlands of Kermanshah 

Province, Western Iran 

Nasrullah Rastegar-Pouyani 1 and Eskandar Rastegar-Pouyani 2 

1 Department of Biology, Faculty of Science, Razi University, Kermanshah, Iran, e-mail: nasrullah® razi.ac.ir. 
'Department of Biology, Faculty of Science, Sabzevar University, Sabzevar, Iran. 

Abstract.- A new and distinctive species of the genus and subgenus Eremias is described from the highlands of 
Kermanshah Province, western Iran at about 1800 m elevation. It is easily distinguished from all other species of 
the typical subgenus (E. velox, E. persica, E. strauchi, E. nigrolateralis, E. lalezharica, E. afghanistanica, E. 
regeli, E. suphani, and E. nikolskii) by a variable number of postmentals (4-5 pairs), smaller size, and distinctive 
color pattern. Furthermore, it can be distinguished by having a combination of characters against any of the 
species in the typical subgenus. Systematics of the genus and subgenus Eremias is shortly discussed. 

Key words.- Lacertidae, Eremias, Eremias (Eremias) montanus. Western Iran , Zagros Mountains, Kermanshah 
province, Siah-Darreh 




Figure 1. Location of Kermanshah province on the Ira- 
nian Plateau. 

Introduction 

The lacertid genus Eremias Fitzinger, 1834 encom- 
passes about 33 species of mostly sand, steppe, and 
desert dweller lizards which are distributed from 
northern China, Mongolia, Korea, Central and south- 
west Asia to southeastern Europe (Rastegar-Pouyani 
and Nilson, 1997). The genus is Central Asian in its 
relationships and affinities (Szczerbak, 1974). About 
15 species of the genus Eremias occur on the Iranian 
Plateau mostly in northern, central, and eastern 
regions (Rastegar-Pouyani and Nilson, 1997; Ander- 
son, 1999). To date, no comprehensive study has been 
carried out on Eremias fauna of the Iranian Plateau 
and the systematic status of most taxa is in great need 
of a revisionary work. Szczerbak (1974), however, 



45 




46 .47 


48 




Iraq 


\_ Iran 








KERMANSHAH ■ 
J PROVINCE Ty 

locality 


CT Hamedan 






□ 
Kermanshah^ 


Lorestan"^_^ 








50 km 



35 



34 



Figure 2. The type locality of Eremias (Eremias) mon- 
tanus, vicinity of the Village of Siah-Darreh, about 
60km northeast of the city of Kermanshah, Kerman- 
shah Province, western Iran. 

revised Eremias and divided it into two distinguished 
genera based on morphological characters: Mesalina 
Gray and Eremias Fitzinger (see under systematic dis- 
cussion). 

As the first record of Eremias, in this paper we 
describe and introduce a new species of this genus 
from the upland and mountainous regions of Kerman- 
shah province, western Iran at about 1800 m eleva- 
tion. This province is located on the western periphery 
of the Iranian Plateau (Fig. 1 ) and a major part of it is 
covered by the Zagros Mountains. The type locality of 
Eremias (Eremias) montanus (sp. nov.) is located in 
an upland area surrounded by the Zagros Mountains 



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Asiatic Herpetological Research 



2001 



with steppe vegetation (e.g., Astragalus, Euphorbia, 
Zygophyllurn), about 60 km northeast of city of Ker- 
manshah, Kermanshah province, western Iran (Fig. 
2). 

Eremias montanus, new species 
(Figs. 3-4) 

Holotype and type locality: An adult female, Field 
number P198, collected by the senior author on 14 
August 1995 from the upland regions of the Zagros 
Mountains, 60 km northeast of city of Kermanshah 
(47° 5'E, 34° 52'N), Kermanshah Province, western 
Iran , at about 1800 m elevation 
Paratypes: Two adult females. Field number PI 99- 
P200, other information as for the holotype. 
Diagnosis and comparison: A small-sized lacertid, 
maximum snout-vent length (SVL)= 57.2mm, tail 
length = 95mm, with 13-14 longitudinal and 27-28 
transverse rows of ventral plates, slightly converging 
posteriorly; with 63-67 small, granular scales across 




middle of dorsum. A species belonging to the subge- 
nus Eremias: subocular reaches mouth edge; one fron- 
tonasal; two supraoculars which are not completely 
separated from frontal and frontoparietals; distance 
between the femoral pores being narrow; color pattern 
"striped"; inhabitant of steppe and mountain-steppe 
landscapes (Szczerbak, 1974: 83). 

On the other hand, it differs from all other species 
of its relevant subgenus based on having several dis- 
tinguishing characters; the color pattern is distinctive 
and it is distinguishable from all other species in this 
character i. e., dorsum uniformly dark-brown without 
spots and ocelli, interrupted by five light longitudinal 
stripes; the vertebral stripe bifurcating on the nape, 
two paravertebrals on each side; a broad dorso-lateral 
stripe containing one or two regularly arranged light 
spots (different from the other Eremias species in this 
character); Furthermore, it differs from each species 
of the typical subgenus in the following character 
combinations (Bischoff and Bohme, 1980; Bohme 
and Szczerbak, 1991; Rastegar-Pouyani and Nilson, 
1997; Szczerbak, 1974) : 

From Eremias nigrolateralis Rastegar-Pouyani 
and Nilson, 1997 in having a much smaller size, lack 
of separation of the third pair of submaxillary shields 
by granular scales (0% versus 100%), lower count of 
gulars (23-24 versus 41-42), variable number of sub- 
maxillary shields (33% versus 0%), reach of femoral 




Figure 3. Eremias (Eremias) montanus holotype. 



Figure 4. Eremias (Eremias) montanus paratypes. 



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Asiatic Herpetological Research 



Vol. 9, p. 109 



pores to the knee (100% versus 0%), and distinct dif- 
ferences in color pattern. 

From Eremias persica Blanford, 1874 in having a 
smaller size, lower count of gulars (23-24 versus 28- 
38), the absence of distinctly keeled upper caudal 
scales (100% versus 75%), variable number of sub- 
maxillary shields (33% versus 4%) and distinct differ- 
ences in color pattern. 

From E. velox (Pallas, 1771) in having a smaller 
size, in the absence of distinctly keeled upper caudal 
scales (100% versus 0%), variable number of sub- 
maxillary shields (33% versus 5%) and in color pat- 
tern. 

From E. strauchi Kessler, 1878 in having a 
smaller size, the absence of distinctly keeled upper 
caudal scales (100% versus almost 0%), variable 
number of submaxillary shields (33% versus 9%) and 
in color pattern. 

From E. lalezharica Moravec, 1994 in having 
variable number of submaxillary shields (33% versus 
0%), higher count of dorsals (63-68 versus 54-59), no 
contact of gulars with the second pair of submaxillary 
shields, lack of a small scale between prefrontals, 
and distinct differences in color pattern. 

From E. afghanistanica Bohme and Szczerbak, 
1991 in a much higher count of dorsal scales (63-68 
versus 44-46), lower count of gulars (23-24 versus 25- 
28), variable number of submaxillary shields (33% 
versus 0%), and in color pattern. 

From E. nikolskii Bedriaga, 1905 in a higher 
count of dorsals (63-68 versus 45-59), variable num- 
ber of submaxillary shields (33% versus 8%), and in 
color pattern. 

From E, regeli Bedriaga, 1905 in a higher count 
of dorsals (63-68 versus 43-61), higher number of 
scales in the 9th- 10th caudal annulus (27-28 versus 
17-25), the absence of distinctly keeled upper caudal 
scales (100% versus 0%), variable number of sub- 
maxillary shields (33% versus 3%), and in color pat- 
tern. 

Description of holotype: An adult female, preserved 
in 75% ethyl alcohol in good condition; body small 
and moderately depressed; a species of the subgenus 
Eremias (Szczerbak, 1974:83); five pairs of submaxil- 
lary shields, first three pairs in contact, the fourth and 
fifth pairs widely separated; first pair of submaxillary 
shileds as large as the fifth and in contact with mental 
anteriorly, with first and second infralabials laterally; 
the fifth sabmaxillary pair each in contact with the 
fourth pair anteriorly, being surrounded by 8 granular 
scales laterally and posteriorly; 7-8 supralabials, 4-5 
of which anterior to subocular which borders the 



mouth; two large nasals in contact with rostral anteri- 
orly, with first and second supralabials laterally, and 
with frontonasal and first loreal posteriorly, the former 
being single, broader than long and laterally in contact 
with first loreal and posteriorly with prefrontals; two 
prefrontals each smaller than frontonasal and almost 
as long as broad and laterally in contact with second 
loreal and posteriorly with frontal and granules of 
supraocular region; only frontonasal with distinct 
concavity; frontal almost as long as prefrontal and 
frontonasal together, broadened and posteriorly and 
laterally partly in contact with supraoculars (and 
partly separated from the latter by 2-3 large scales, not 
by granules) and posteriorly with frontoparietals; two 
frontopariatals almost as large as a single supraocular, 
laterally being in contact with the second supraocular, 
and posteriorly with interparietal and parietals, the 
former being small and relatively lozenge-shaped, 
surrounded by frontoparietals and parietals; two vey 
large and plate-like parietals, almost as long as broad, 
being in contact behind interparietal; a vestigial 
occipital; two loreals, first one small, surrounded by 
first two supralabials, nasal, frontonasal, and the sec- 
ond loreal which is distinctly large with an evident 
keel; 6-6 supraciliaries, separated from supraocular by 
a series of 42-44 granules; postocular elongate, sur- 
rounded by granules anteriorly; temporal region- 
mostly covered by granular scales becoming large 
towards the orbit, more than 100 on each side; tym- 
panic scale distinct and elongated obliquely, almost 
the same size on both sides; tympanum vertically 
elongated, slightly larger than orbit; no distinct 
supratemporal; subocular huge, broader than long 
with a distinct ridge being extensively in contact with 
the lower edge of orbit; lower eyelids with a semi- 
translucent membrane made up of about 22 enlarged 
scales; collar well pronounced , not serratted, made up 
of 10 scales, the two medial ones the largest; gular 
fold weakly developed, 23-24 gulars from symphysis 
of chin shields to median gular, becoming enlarged 
posteriorly; 13-14 longitudinal and 27-28 transverse 
rows of almost squarish ventral plates from collar to 
hindlimbs; anterior series of ventrals to some extent 
irregular, median ventral longer than broad; dorsal 
scales juxtaposed, smooth, granular, becoming slighly 
larger posteriorly, 63-68 scales across the middle of 
dorsum, and about 160-164 scales in a single row 
from occiput to a point just above the vent; proximal 
caudals larger than posterior dorsals but the change 
being gradual; caudals becoming large, elongate, and 
slightly keeled distally, arranged in distinct whorls, 
27-28 scales in the 10th whorl behind the vent; upper 
forearm covered dorsally by enlarged, juxtaposed, and 
almost lozenge-shaped scales; lower forearm covered 



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Asiatic Herpetological Research 



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with granules; upper hindlimbs covered externally by 
granules, externally by large shields; tibia covered 
dorsally by slightly pointed granules, ventrally by 
large plates which are slightly keeled, two plates in a 
transverse row; no fringes on the toes, 18-20 uni-and 
bi-carinate lamellae under fourth toe, proximal part of 
lower fourth toe containing two complete rows of 
lamellae, distal part with a single row (in this charac- 
ter it is quite different from all other species of its rel- 
evant subgenus); 18-19 femoral pores in each side, the 
two series separated anteriorly by a narrow space con- 
sisting of three scales; preanal region encompassing 
24 large shields, the four median ones being the larg- 
est; 6 plates in longitudinal row from the space 
between femoral pores to anterior edge of the vent. 

Coloration and color pattern: Dorsum uniformly 
dark-brown without spots and ocelli, interrupted by 
five light, narrow longitudinal stripes: one vertebral 
bifurcating on the nape, two paravertebrals on each 
side; a broad dorso-lateral stripe containing one or 
two regularly arranged light spots; a ventro-lateral 
series of dark-brown spots, to some extent forming a 
stripe; upper surface of head uniformly olivish-brown; 
temporal and labial regions suffused by dark-brown; 
submaxillary region light-gray-cream; ventral region 
dirty white, suffused by bluish-brown; upper surface 
of limbs dark-brown containing numerous light spots; 
proximal upper surface of tail brown, distal part light 
brownish-gray; lower surfaces of limbs and tail yel- 
lowish-white. 

Description of paratypes: The paratypes, two adult 
specimens, one male (PI 99) and one female (P200) 
approximate the holotype in almost all pertinent 
details. However, there are some minor differences 
between paratypes and the holotype as follows: 

Male paratype: five submaxillary shields on the right 
side but only four on the left, the fifth vestigial; 13-14 
longitudinal and 30-31 transverse ventral plates; 10- 
1 1 collars, 3-4 median ones the largest; 23-24 gulars 
in a longitudinal row from chin shields to collar; 20- 
20 femoral pores, separated by three small scales; 23- 
24 lamellae under fourth toe; 23-27 scales around 
10th tail annulus; 8-9 labials, 5 of them anterior to 
subocular; 6-6 lower labials; lower nasal resting on 
the first two supralabials; temporal scale vestigial; 62- 
63 scales around widest part of dorsum; 162-167 
scales in a single longitudinal row from occiput to 
vent. 

Measurements (mm): SVL = 58.5; TL = incomplete; 
Forelimb = 24; Hindlimb = 39; HL = 15.5. 
Female paratype: only four pairs of submaxillary 
shield, the fifth vestigial; 15-17 longitudinal and 31- 



33 transverse rows of ventral plates; 9 collars, 4-5 
median ones the largest; 24-25 gulars in a single lon- 
gitudinal row from chin shields to collar; 20-20 femo- 
ral pores, separated by a space corresponding to two 
scales; 25-26 scales on the 10th tail annulus, 24-25 
subdigital lamellae under fourth toe; 8-9 supra- and 4- 
6 infralabials; temporal scales relatively small; frontal 
separated from the first supraocular by two large 
scales (not by granules); 65-67 scales around widest 
part of dorsum; 163-167 scales in a single longitudi- 
nal row from occiput to vent. 

Measurements (mm): SVL - 52.9; TL = incomplete; 
Forelimb = 23; Hindlimb = 35.5; HL = 13.6. 

In color pattern they are quite similar to the holotype. 

Habitat: During field work on the western regions of 
the Iranian Plateau in 1995, we surveyed the Zagros 
Mountains and the nearby mountain steppes in the 
northern parts of Kermanshah province. 60 km north- 
east of Kermanshah city, in the highland steppes, in 
vicinity of the Village of Siah-Darreh in an area 
named Sarpal, we came across to three specimens of a 
new taxon of the genus Eremias, described here as a 
new species. The habitat, which is surrounded by the 
Zagros Mountains, is characterized by a mountain- 
steppe; the vegetation is luxuriant steppe association: 
mainly Astragalus, Euphorbia, Artemisia, and 
Amigdalus as well as various species of the families 
Graminaceae and Compositeae (Fig. 5). 

Since it is a mountainous region with relatively 
high elevation, the winter being harsh and cold, the 
summer being mild and rather short. All the speci- 
mens were foraging around the shrubs probably look- 
ing for prey. They were quite shy and wary and very 
difficult to capture. When alarmed, they took refuge 
under the shrubs or inside the underground holes. One 
of the most effective anti-predatory adaptations 
evolved in these lizards is the ability to lose the tail 
(autotomy) when being touched by the predators (or 
collectors). Hence, we could only collect one speci- 
men with a complete tail and the other specimens lost 
their tails during capturing. 

In September 1998, we re-surveyed the type local- 
ity in order to find more specimens of this species but 
without success. Whether it being a relictual and rare 
species, confined only to the type locality, or being 
distributed over a wider area in the western margin of 
the Iranian Plateau is yet to be established. 

With regard to the occurrence of Eremias monta- 
nus in the highlands of Kermanshah province, the 
Procter record of E. velox, as the westernmost record, 
from Kuretu (Iran-Iraq border) (Procter, 1921:252) 
should seriously be reconsidered. 



Asiatic Herpetological Research 



Vol. 9. p. I 1 1 



<# 



js»* h »- 



"ym 



-*> 



•" E 



ilfc 



•r 



■w 





■-** 



Figure 5. Habitat and type locality of Eremias (Eremias) montanus. 60 km northeast of Kermanshah, vicinity of 
Siah-Darreh village, Kermanshah Province, Western Iran. 



Etymology: Eremias montanus is so named as it is 
apparently restricted in distribution to the upland and 
mountainous steppes of northeastern regions of Ker- 
manshah province, western Iran. 

Taxonomic account: As mentioned before, so far, the 
most complete work done on the complicated genus 
of Eremias (sensu lato) is of Szczerbak (1974) who 
studied almost all species and species complexes of 
this genus throughout the range. Based on morpholog- 
ical characters and geographic distribution, S/.czerbak 
(1974) subdivided the inclusive genus Eremias (s. 1.) 
into two distinct genera; the genus Mesalina as a 
north African and lowland southwest Asian clade, and 
the genus Eremias (sensu stricto) which is mainly 
occurring in Central and northeast Asia. (Szczerbak. 
1974). 

Furthermore. Szczerbak (1974) subdivided Ere- 
mias (s.s) into five distinct subgenera: Eremias (Szcz- 
erbak, 1974: 83), Rhabderemias (Szczerbak, 1974: 
201 ), Ommateremias (Szczerbak, 1974: 146). Parere- 
mias (Szczerbak, 1974: 22-23), and Scapteria ( Szcz- 
erbak, 1974:247). 

Except for the subgenus Pareremias, which is a 
Central and east Asian clade, all of the major species 
groups of the genus are presented on the Iranian Pla- 



teau (Anderson, 1999). Arnold ( 1986) who studied the 
hemipenes of lacertids supported the Szczerbak's sub- 
generic names. In a more recent study Arnold placed 
Eremias as the sister (axon of a clade including Acan- 
thodactylus, Mesalina, and Ophisops-Cabrita 
(Arnold. 1989:238, 240 ). But Mayer and Benir 
( 1994) have proposed a different scenario. According 
to these authors, Eremias is the sister taxon of 
Mesalina and both of them belong to a larger clade 
also containing Omanosaura and Ophisops. They 
believe that Eremias is not closely related to Acantho- 
dactylus. 

The Czech Biological Expedition to Iran in 1996 
collected 8 specimens of an undetermined species of 
Eremias related to E. persica from the Zagros Moun- 
tains in Esfahan province at about 2000-2200 m ele- 
vation (Frynta et al., 1997: 9-10). Whether it is a new 
taxon or just a variety off. persica is yet to be known. 

Material examined: Eremias montanus (n = 3): P 
198-200 (Field number), from around the Siah-Darreh 
Village (about 1800 m elevation). 60 km northeast 
city of Kermanshah. Kermanshah province, western 
Iran. 



Vol. 9. p. l : 



Asiatic Herpetological Research 



2001 



Eremias nigrolateralis (n = 2): GNHM. Re. ex. 
5147-5148, from 150 km northeast of Shiraz. Fars 
province, south-central Iran. 

Eremias persica (n = 4): GNHM. Re. ex. 5159- 
5162. from 150 km northeast of Shiraz. Fars province, 
south-central Iran. 

Eremias persica (n = 28) : GNHM. Re. ex. 5163- 
5190, from 45 km east of Arak on the road to Qum, 
Markazi province, north-central Iran. 

Eremias persica (n = 4) : GNHM. Re. ex. 5191 - 
5194, from 65 km west of Tehran, between Eshtehard- 
Saveh, Tehran province, northern Iran. 

Eremias persica (n = 2) : GNHM. Re. ex. 5195- 
5196. from 45 km east of Golpaygan, Esfahan prov- 
ince, central Iran. 

Eremias persica (n = 4) : GNHM. Re. ex. 5197- 
5200. from 50 km north of Delijan on the road to 
Qum. Markazi province, north-central Iran. 

Eremias persica (n = 1) : GNHM. Re. ex. 5201, 
from 50 km north of Abadeh. Fars province, south- 
central Iran. 

Eremias persica (n = 1) : GNHM. Re. ex. 5202. 
from 50 km east of Hamedan on the road to Qazvin, 
Hamedan province, western Iran. 

Eremias persica (n = 3) : GNHM. Re. ex. 5203- 
5205. from 5 km west of Takestan on the road to Zan- 
jan, Zanjan province, northwestern Iran. 

Eremias velox (n = 4) : GNHM. Re. ex. 5122(1- 
4). from around the Carin River. 250 km E-SE Almaty 
(Alma Ata). Kazakhstan. 

Eremias velox (n = 2) : GNHM. Re. ex. 5120(1- 
2), from Mulali Kurozek. eastern Kazakhstan. 

Eremias velox ( n = 2 ) : GNHM. Re. ex. 5121(1- 
2), from the Taldi Korgau District, northeast Kazakh- 
stan. 

Eremias velox (n = 2) : GNHM. GK. 18881 (1-2). 
from Archenjan Village ( 1 ), and 30 km north of Mary 
(2), Turkmenistan. 

Eremias strauchi (n = 3) : GNHM. Re. ex. 441 1 
(1-3), from Golestan National Park, Mazandaran 
province, northeastern Iran. 

Abbreviations : GNHM. Re. ex. = Gothenburg Natu- 
ral History Museum. Reptilia exotica; GNHM. GK. = 
Gothenburg Natural History Museum. General Kata- 
logue. 

Acknowledgements 

We wish to thank the Razi University authorities (Ker- 
manshah-Iran) for their generous help and support 
during field work in western Iran. 



We thank the Gothenburg Natural History Museum 
(Gothenburg-Sweden) for loan of Eremias specimens. 
Also we thank Dr. Michael Golubev for translation of 
the relevant Russian literature. 

Literature Cited 

Anderson, S. C. 1999. The Lizards of Iran. Society for 
the study of Amphibians and Reptiles 442 pp. 

Arnold, E. N. 1986. The hemipenis of lacertid lizards 
(Sauria : Lacertidae): structure, variation and system- 
atic implications. Journal of Natural History 20:1221- 
1257. 

Arnold, E. N. 1989. Towards a phylogeny and bioge- 
ography of the Lacertidae : relationships within an 
Old- World family of lizards derived from morphol- 
ogy. Bulletin of the British Museum (Natural His- 
tory). Zoology 55(2):209-257. 

Bischoff, W. and W. Bohme. 1980. Der systematische 
Status der turkischen Wustenrenner des Subgenus 
Eremias (Sauria: Lacertidae). Bonner Zoolgische 
Beitrage, N. F. 26:297-306. 

Bohme. W. and N. N. Szczerbak. 1991. Ein neuer 
Wustenrenner aus dem Hochland Afghanistans. Ere- 
mias (Eremias ) afghanistanica sp. n. (Reptilia : Sau- 
ria : Lacertidae). Bonn. Zool. Beitr. 42:137-141. 

Frynta. D.. J. Moravec, J. Cihakova, J. Sadlo, Z. Hod- 
kova, M. Kaften. P. Kodym. D. Krai. V. Pitule. and L. 
Sejna. 1997. Results of the Czech Biological Expedi- 
tion to Iran. Part 1. Notes on the distribution of 
amphibians and reptiles Acta Societatis Zoologicae 
Bohemicae 61 :3-17. 

Mayer. W. and G. Benyr. 1994. Albumin-Evolution 
und Phylogenese in der Familie Lacertidae (Reptilia: 
Sauria). Annalen Naturhistorischen Museums in Wien 
96B:62 1-648. 

Moravec. J. 1994. A new lizard from Iran. Eremias 
(Eremias ) lalezharica sp. n. (Reptilia: Lacertilia: 
Lacertidae). Bonner Zoolgische Beitrage 45(1 ):6 1-66. 

Procter. J. B. 1921. Further lizards and snakes from 
Persia and Mesopotamia. Journal of the Bombay Nat- 
ural History Society 28(1 ):25 1-253 

Rastegar-Pouyani. N. and G. Nilson. 1997. A New 
Species of Eremias (Sauria: Lacertidae) from Fars 
Province, South-Central Iran. Russian Journal of Her- 
petology4(2):94-101. 

Szczerbak, N. N. 1974. Yashchurki Palearktiki (Pale- 
arctic species of Eremias ). Kiev. 295 pp. 



2001 



Asiatic Herpetological Research 



Vol.9, pp. 113-121 



Lizards of the Northern Mongolian Deserts: Densities and Community 

Structure 

KONSTANTIN A. ROGOVIN 1 , DMITRY V. SEMENOV 1 , AND GEORGY I. SHENBROT 2 

'A.N.Severtzov Institute of Ecology and Evolution, Russian Academy of Science, Leninsky pr. 33 Moscow 

1 17071, Russia, ~ Ramon Science Center, Ben-Gurion University of the Negev, P.O. Box 194, Mizpe Ramon, 

80600, Israel 

Abstract.- Spatial organization and population densities of three-species lizard community was studied in the 
Gobi Desert, Mongolia. To evaluate the effect of habitat variables on the distribution and abundance of each 
species we used the stepwise procedure of factor selection with ANOVA on each step. To describe the 
distribution of species' spatial niches in the space of environmental variables, we used stepwise discriminant 
function analysis (DFA). The number of species in 1-ha grid areas varied from to 4. Phrynocephalus versicolor 
was the only species distributed over the 91% of grids occupied. There was a positive relationship between 
distribution and local species abundance. A set of two to three habitat variables determined the abundance of each 
species. The result of DFA signify to the well pronounced segregation, but not even distribution of species spatial 
niches in the space of resources. 



Key words.- Lizards, Mongolia, community, ecology, density 




91* 96* 101* 106° 

Fig 1. Map of Mongolia and location of sites where data were collected. 
Eastern Gobi, 4-Barun-Churay Basin. 



111* 116' 

Southern Gobi, 2-Western Gobi, 3- 



Introduction 

The reptile communities of Mongolian deserts are 
characterized by several specific features discussed 
elsewhere (Ananjeva and Semenov, 1986; Borkin and 
Semenov, 1984; Munkhbajar, 1976; Semenov and 
Borkin, 1986; Semenov and Shenbrot, 1988). There 
are few species in the fauna with low species richness 



at sites, a low level of species turnover between habi- 
tats, low abundance of most species and high domi- 
nance of only one species, Phrynocephalus versicolor. 
A few common species have rather broad spatial 
niches, diverse behavioral and physiological charac- 
teristics (diverse range of thermobiological patterns, 
wide active search for food items, etc.). 



Vol. 9, p. 114 



Asiatic Herpetological Research 



2001 



The degree of interest in comparative studies of 
structure and function of reptile communities 
increased dramatically after seminal papers written by 
E. Pianka (Pianka, 1973, 1975). Most of the ensuing 
studies were devoted to the species rich and diverse 
communities of desert lizards in Australia, south- 
western North America and southern Africa (Case, 
1983; Fuentes, 1976; Henle, 1989; Inger, Colwell, 
1977; Millado et al., 1975; Pianka,1986; Scheibe, 
1987; Shenbrot et al., 1992; Simbotwe, 1984). At the 
same time study of species poor communities in the 
Central Asian desert can provide a significant infor- 
mation not only in comparison with other continents, 
but also can help us to understand better which factors 
rule in reality structure and dynamic of lizard commu- 
nities of many species. Up to now there were only two 
examples of such studies made in China (Chang et al., 
1993;Luietal.,1992). 

The main objective of this paper was to study the 
specific features of spatial organization and popula- 
tion densities of three-species lizard community in the 
Gobi Desert, Mongolia. The study was conducted dur- 
ing a long-term research program on the biodiversity 
of the Mongolian desert biota, and was sponsored by 
the Permanent Soviet-Mongolian Biological Expedi- 
tion. 

Material and Methods 

Mongolian desert 

A map (Fig. 1) illustrates the location of desert 
regions of Mongolia. Three desert regions to the south 
of the Altai Mountains are partly separated from one 
another by chains of low mountains and hills. These 
three are Trans-Altai Gobi (South), Alashan Gobi 
(East) and Sungarian Gobi or Barun-Churay Basin 
(West) (Yunatov, 1950). Besides these deserts there 
are desert areas between the Altai and Hangai moun- 
tains, usually called Western cold deserts, and some 
arid lands in the Great Lakes and Ubsu-Nur Basins 
(northwest). 

Three arid subzones of the Mongolian desert are 
defined (Sokolov and Gunin, 1986): extra — arid 
desert (<50 mm of rainfall per year), real desert (50— 
100 mm per year) and steppe — like desert (100—150 
mm per year). However, the climatic border that 
restricts the distribution of plants (Kazantseva, 1986; 
Volkova et al., 1986) and animals (Podtyazhkin and 
Orlov, 1986; Semenov and Borkin, 1986) exists only 
between the southern part of Trans- Altai Gobi (< 50 
mm per year) and the northern waste belt of desert 
lands with more predictable precipitation (100-200 
mm per year). The narrow real desert subzone appears 



transitive with no specific features of vegetation. 
Thus, only the southern (extra-arid) and northern sub- 
zones are well pronounced. 

The southern desert occurs mostly in the Trans- 
Altai Gobi and is characterized by a few very dry, 
unproductive biotopes inhabited by five lizard species 
(two agamids, two gekkonids, and one Eremias spe- 
cies). Among this group only one species, Phryno- 
cephalus versicolor is common in the northern 
subzone (Semenov and Borkin, 1986). 

The northern deserts, which extend to the south- 
west, west, and south-east of Mongolia and along 
both slopes of the Mongolian Altai and Gobi-Altai 
mountains, are characterized by pronounced microre- 
lief and rich vegetation, although the main landscape 
types are the same as in the southern subzone. The 
vegetation in rock and gravel valleys consists of 
perennial grass (Stipa), forbs, onions and succulents, 
and a variety of annual plant species. Shrub vegetation 
is often associated with foothills and sand dunes, or is 
spread along the dry river beds (Lavrenko, 1978). The 
difference in climate between western and eastern 
parts of the northern Mongolian desert is not pro- 
nounced (Murzaev, 1952); some differences exists in 
the composition of the flora (Yunatov, 1950). 

Lizard species 

There are four lizard species inhabiting northern 
Mongolian deserts: Alsophylax pipiens, Phrynoceph- 
alus versicolor, Eremias przewalskii and E. multiocel- 
lata. Among these, only the three last mentioned 
species are abundant and relatively widespread. 

Data collection 

We collected data during three field trips to the Mon- 
golian northern deserts in June-August 1985, 1986 
and 1988. Forty five 1-ha grids were established in the 
Northern desert subzone (see map, Fig 1 ). Grids were 
distributed so that they covered the whole range of 
habitats from the middle slopes of the mountains to 
the clay basins and sand dunes. Each habitat type was 
sampled equally. Two factors determined the number 
of grids at a desert region: diversity of habitat types 
and abundance of lizards. Each grid was divided into 
25 smaller sample plots, 20 x 20 m. The centers of 
sample plots were marked with 50 cm aluminum 
stakes. Lizards were sampled by repeated, regular 
search of established grids during two to four consec- 
utive days during periods of their maximal diurnal 
activity. Nearly all encountered lizards were captured 
by hand. Most of the surveys accounting were con- 
ducted before the appearance of hatchlings. In the rest 
of the cases hatchlings were not counted. Each cap- 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 115 



Table 1. Density (no. ha ) of lizard species. 



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/? versicolor 



E przewalskii 



E. multiocellata 



1.7±0.9 52.8+8.9 54.4±8.7 36.6±8.2 5.5±4.5 41.2+6.3 0.8±0.4 

0-3 24-91 32-83 10-106 1-10 22-64 0-3 

2.6±1.3 16.3±6.3 0.9±0.5 

0-18 0-44 0-4 

2.1+1.3 0.4±0.4 0.5±0.4 2.0±1.8 
0-12 0-2 0-5 0-11 



tured lizard was marked by toe-clipping (the most dis- 
tal phalange only) and/or by a number painted with 
permanent color marker on its back, and released. 
Densities of lizard species were estimated by the total 
number of animals caught on a grid. The reliability of 
density estimates was verified by recapture of marked 
individuals and by the observation of marked lizards 
on grids. Two days were sufficient to catch > 80% of 
lizards. 

A 0.5 kg soil sample was taken from the center of 
each plot for laboratory texture analysis. The number 
of shrubs (by species) in each plot was counted in a 
circle, 5 m in radius. To determine vegetation cover 
and volume by height layers within each plot, the 
height and diameter of crown of the shrubs (up to 30 
shrubs of each species in the grid) were measured. 
The abundance of annuals was evaluated by clipping 

all annuals on 0.25 m" sample plots (4 sample plots 
placed at random in each of 25 plots in 1-ha grid 
area). Twenty three parameters were used in the sub- 
sequent analysis (Table 2). Data on 1 125 descriptions 
of sample plots and 1710 records of 3 lizard species 
were used in the analysis. In total, we recorded 1528 
individuals of the agamid lizard P. versicolor, 142 of 
the lacertid lizard Eremias przewalskii and 40 E. mul- 
tiocellata. 



Statistical data processing 

To evaluate the effect of habitat variables on the distri- 
bution and abundance of each species we used the 
stepwise procedure of factor selection (Shenbrot, 
1988) with ANOVA of each step. Before analysis, the 
original values of species' densities and habitat vari- 
ables were transformed to an interval scale with five 
intervals for habitat variables and three intervals for 
species' abundance variables. The influence of each 
variable was estimated separately by ANOVA and the 
variables with nonsignificant effect were omitted. In 
the next step, each of variables that was not omitted 
earlier was added separately to the most powerful 
variable to select the most powerful pair of variables. 
This procedure was repeated until all possible vari- 
ables were included in the analysis or until the addi- 
tion of the new variable does not increase the 
proportion of explained dispersion. 

To describe the distribution of species' spatial 
niches in the space of environmental variables (eco- 
logical space) and to reduce the dimensionality of this 
space, we used stepwise discriminant analysis. Dis- 
criminant axes were calculated based on the data set 
consisting of the values of habitat variables for each 
point of lizard's registrations and with lizard species 
as a grouping variable. 



Vol. 9, p. 116 



Asiatic Herpetological Research 



2001 



Results 

Densities and distribution through habitat 
types 

The results of lizard density and diversity estimations 
on the 1-ha grids indicate the low local species diver- 
sity in Mongolian deserts (Table 1). The number of 
species in our grid areas varied from zero to three. 
Among 45 grids there were two grids with no lizards, 
21 grids with only one species. 18 grids with two spe- 
cies and four grids with three species. Phrynoceph- 
alus versicolor was the only species distributed over 
the most number of grids (91% of grids occupied). 
Eremias przewalskii was found on 33%, E. multioce- 
lata on 24% and A. pipiens on 4% of grids. Regarding 
distribution through the main habitat types (Table 1), 
P. versicolor was also the most widely distributed spe- 



cies but with the pronounced habitat preference. The 
second was E. multeocellata, and the third was E. 
przewalskii. The last species had the most restricted 
habitat preference, namely sandy-loess hills in saline 
depressions with shrub vegetation of Nitraria sp. 

The abundance of P. versicolor was positively cor- 
related with its broad distribution. The density of this 
species varied from 1 to 106 individuals per 1-ha 
(Table 1). The second most abundant E. przewalskii 
(44 individuals per lha maximum), and the third was 
E. multiocelata (11 individuals per 1-ha maximum). 
All three species coexisted at rather high densities in 
saline depressions with sandy-loess hills covered with 
Nitraria sp. shrubs. There were no pronounced nega- 
tive correlation between the densities of two dominant 
species on grids situated within this habitat type (P. 
versicolor-E. przewalskii: R"=0.04, n=20, ns). 



Table 2. Designation and description for the 23 habitat variables included in the analysis. 



Mnemonic 



Variable 



Unit 



RCK 

GRW 

CLY 

SCS 

SCH 

WDS 

WDD 

NRB 

FRB 

AGR 

ANN 

ALL 

PGR 

MIC 

HAL 

sue 

HLX 

NIT 

SHC 

SV1 

SV2 

SV3 

SV4 



Content of rocks in the soil 

Content of gravel in the soil 

Content of clay in the soil 

Sand cover area 

Sand cover height 

Dry river bed area 

Dry river bed depth 

Number of rodent burrows 

Abundance of annual forbs 

Abundance of annual grasses 

Overall abundance of annual grasses and forbs 

Perennial Allium covet 

Perennial grass cover 

Cover of microphyllous shrubs 

Cover of halophytuos shrubs 

Cover of small succulent shrubs 

Cover of Haloxylon 

Cover of Nitraria 

Overall shrub cover 

Perennial plant crown volume at the level 0-25 cm 

Perennial plant crown volume at the level 25-50 cm 

Perennial plant crown volume at the level 0.5-1 m 

Perennial plant crown volume at the level 1-2 m 



%% 

%% 

%% 

%% 

cm 

%% 

cm 

no/sq.m. 

no/sq.m. 

no/sq.m. 

no/sq.m. 

%% 

%% 

%% 

%% 

%% 

%% 

/o /o 

o/ o/ 
/o /o 

%% 

%% 

0/ O/ 

/o /o 
o/o. 



Vo% 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 117 



1 
0.5 



-0.5 

-1 



AGR 



1 
0.5 


-0.5 

-1 



NIT 



SHV2 



SHV2 



1 

0.5 



-0.5 

-1 




GRV 



1 
0.5 


-0.5 

-1 



1 



Fig 2. Habitat use profiles of P. versicolor. For each 
habitat variable rank, mean use by species is plotted 
as a difference between capture frequency on plots of 
this rank and capture frequency of all plots. The size of 
histograms therefore denote the selectivity (positive or 
negative) of habitat use. For mnemonics see Table 2 

Spatial niches 

There was a statistically significant influence of habi- 
tat variables on the distribution and abundance of all 
three studied lizard species (Table 3). For each of 
these species we extracted a set of two to three habitat 
variables, determining 6.6-28.7% of observed vari- 
ance in abundance. Densities of two species (P. versi- 
color, E. przewalskii) were moderately affected by 
habitat variables, whereas density of E. multiocellata 
was weakly affected. 

There was a statistically significant influence of 
habitat variables on the distribution and abundance of 
all three studied lizard species (Table 3). For each of 
these species we extracted a set of two to three habitat 
variables, determining 6.6-28.7% of observed vari- 
ance in abundance. Densities of two species (P. versi- 
color, E. przewalskii) were moderately affected by 
habitat variables, whereas density of E. multiocellata 
was weakly affected. 




CLY 



1 

0.5 





-0.5 
-1 



Fig. 3. Habitat use profiles of E przewalskii. For expla- 
nation see Fig. 2. 

FRB 




MIC 




Fig. 4. Habitat use profiles of E multiocellata. For 
explanation see Fig.2. 

Patterns of habitat usage based on selected vari- 
able sets for each species are presented on Figures 
2-4. Phrynocephalus versicolor clearly avoided 
microsites with low gravel content, very low and very 
high annual grass abundance, moderate and high 
shrub crown volume in the level 0.25-0.5 m and pre- 
ferred microsites with moderate gravel content, mod- 
erate annual grass abundance and very low shrub 



Vol. 9, p. 118 



Asiatic Herpetological Research 



2001 



Table 3. Summary of ANOVA analyses of influence of habitat variables on individual species' abundance. Values 
are proportions of total dispersion determined by given variable. Total proportion of variance determined by a set 
of variables may be greater than sum of influences of individual variables as a result of high-order interactions. 



Species V a r 


i a b 1 e s 






Total 


GRV CLY AGR 


FRB MIC 


NIT 


SHV2 




P. versicolor 0.0226 0.0387 






0.0458 


0.1900 


E. przewalskii 0.0593 




0.1592 


0.0102 


0.2872 


E multeocellata 


0.0180 0.0232 






0.0662 


crown volume in the level 0.25-0.5 m (Fig. 2). Ere- 


Discussion 









mias przewalskii avoided microsites with high and 
very high clay content, very low Nitraria cower, very 
low shrub crown volume in the level 0.25-0.5 m, and 
preferred microsites with low clay content, moderate 
to high Nitraria cower, moderate to high shrub crown 
volume in the level 0.25-0.5 m (Fig. 3). Eremias mul- 
tiocellata avoided microsites with very high and very 
low forb abundance and microphyllous shrub cover, 
and preferred microsites with moderate forb abun- 
dance and rather high microphyllous shrub cover (Fig. 
4). 

Structure of ecological space occupied 

The results of reducing habitat space dimensionality 
using discriminant analysis showed that division of 
this space by lizard species occurred along the first 
two axes. Both axes accounted in sum for 100 % of 
variance and reflected some complex environmental 
gradients (Table 4). The first axis represented a gen- 
eral gradient of decreasing rock and gravel content in 
the soil as well as an increase in the sand cover area, 
sand mound height, number of rodent burrows and 
general shrub (especially Nitraria) cover. This axis 
characterized the between-habitat component of envi- 
ronmental variation rather than within-habitat varia- 
tion, and described in general spatial segregation of 
lizard species according to their preference of physi- 
ognomically distinctive habitats. The second axis 
reflected an increase of dry river bed area, Allium 
abundance and microphyllous shrub cover, and char- 
acterized both between- and within-habitat compo- 
nents of species segregation. The first axis described 
habitat division between E. przewalskii and two other 
species, whereas the second axis reflected habitat seg- 
regation between E. multiocellata and two other spe- 
cies (Fig. 5). All three lizard species had spatially 
remote niche centers and less than 30% niche overlap 
(Fig.5). 



Our data confirm the general opinion of low reptile 
species richness in Mongolian deserts. There were 
only four 1-ha grids where three lizard species coex- 
isted. If we take into account rare encounters with 
snake species, the maximum reptile species number is 
five. Another opinion about wide spatial niches of 
Gobian lizards (Semenov and Borkin, 1986) partly 
contradicts our results. At the one hand P. versicolor is 
the an abundant species that can be found everywhere 
in the Gobi desert, including sandy, clay and rocky 
habitats. This feature of its distribution distinguishes 
this species greatly from its western congeners that 
share habitat types in Middle Asia (Shenbrot et al., 
1991 ). At the other hand, P. versicolor did have well 
pronounced spatial preferences as it is seen from 
Table 1. The two species of Eremias lizards had dis- 
tinctly non-preferable habitats. Eremias przewalskii 
was mainly restricted to one habitat type, and E. mul- 
tiocellata to two habitat types. 

Results of analysis of microhabitat preferences 
demonstrate the existence of significantly distinct 
environmental variables determining spatial distribu- 
tion of each lizard species. The variables elucidated 
can be regarded as axes of species spatial niche. Spa- 
tial niches for P. versicolor and E. przewalskii 
appeared to be determined by three axes of environ- 
mental variation and for E. multiocellata by two axes. 
The distribution of species along each axes can be 
interpreted in accordance with the biological charac- 
teristics of each species. The absence of P. versicolor 
at microsites with dense vegetation is explained by 
typical Phrynocephalus locomotion on straightened 
legs and by the group-specific visual orientation when 
foraging (in contrast to Eremias, which look for food 
items using olfaction). Phrynocephalus versicolor 
preferred microsites with moderate gravel content, 
moderate annual grass abundance and very low shrub 
crown volume, which characterizes the species as an 
inhabitant of stony and gravel desert valleys (Fig. 2). 
Eremias przewalskii selected microhabitats in accor- 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 119 



Fig. 5. Seventy-five percent confidence ellipses for the species observations on two discriminant axes (DF1 and 
DF2). Em- Eremias multiocellata, Ep- Eremias przewalskii, Pv - Phrynocephalus vesicolor. 



fa 

Q 




DF1 



dance with food abundance and a species-specific tac- 
tic of antipredator behavior. It preferred microsites 
with moderate to high Nitraria cover and moderate to 
high shrub crown volume at 0.25-0.5 m (Fig. 3). In 
summer E. przewalskii feed predominantly on berries 
and young green twigs of Nitraria, and also find pro- 
tection under the dense cover of the crowns of low 
spiny shrubs. In contrast to E. przewalskii, E. multio- 
cellata avoided microsites with very high annual plant 
abundance and shrub cover, but also preferred micro- 
sites with moderate and rather high values of these 
variables (Fig. 4). This difference can be interpreted 
in accordance with thermobiological and size charac- 
teristics of two Eremias species. The larger species, E. 
przewalskii is not so quick as the smaller E. multiocel- 
lata. The first one digs well in soft soil, and ther- 
moregulates climbing on or escaping under the shrub 
periodically, being active throughout the day. Small E. 
multiocellata that occupy habitats with relatively low 
vegetation cover (with small sparsely-distributed 
shrubs) must cross open sites in search for food items 
and escape predator's attacks by quickly rushing into 
small shrubs or burrows. This species reduces heat by 
escaping into burrow during the day-time. 

Another result of the above comparison is the pos- 
itive relationship between distribution and local abun- 
dance among the species considered. Distribution 



here means not the size of the species range area, but 
the number of sites where each species was found. 
Phrynocephalus versicolor was encountered on 41 
grid areas and had the highest population density 
(mean: 37.3, median: 26 ind/ha, maximum: 106 ind/ 
ha). With the edge-effect correction (Semenov, 1991) 
maximum density was 70.02 ind/ha (175.5 g/ha biom- 
ass). Eremias przewalskii was found on 1 5 grid areas 
and was the second abundant species (mean: 9.5, 
median: 4 ind/ha, maximum: 44 ind/ha). Eremias mul- 
tiocellata was found on 1 1 grid areas and its maxi- 
mum density was 12 ind/ha (mean: 3.6, median: 2 ind/ 
ha). Alsophilax pipiens was met on two grids with 
density 3 ind/ha. 

Brown (1995) explains this rather common rela- 
tionship by the "Hutchinsonian niche model" (see 
also for one species, Brown 1984), suggesting that 
"the species that is slightly more tolerant of some abi- 
otic conditions or biotic interaction or is slightly bet- 
ter able to use some resource should not only be able 
to occur in more places but also to attain higher abun- 
dance in some of those places." In the approach we 
use here niche breadth reflects microhabitat require- 
ments of each species, namely the range of microcon- 
ditions where each lizard species occurred. Diversity 
of these microhabitats is not necessarily correlated 
with the diversity of macrohabitats as well as with the 



Vol. 9, p. 120 



Asiatic Herpetological Research 



2001 



Table 4. Summary of discriminant analysis of the habi- 
tats of lizard species. DF1 and DF2 are the first two 
components (all are significant, P<0.001). Mnemonics 
for habitat variables are from Table 1 . 





DF1 


DF2 


Eigenvalue 


0.351 


0.102 


Chi-square 


676.1 


165.5 


Cumulative % of variance 


62.58 


37.42 


Factor loading 






RCK 


-0.350 


0.295 


GRV 


-0.291 


0.124 


SCS 


0.544 


-0.221 


WDD 


-0.091 


0.241 


NRB 


0.442 


0.253 


SHH 


0.627 


0.190 


ALL 


-0.141 


0.252 


MIC 


-0.021 


0.410 


NIT 


0.642 


0.193 


SHC 


0.650 


0.148 


SV1 


0.712 


0.067 


SV2 


0.693 


-0.069 



size of the landmass studied. Phrynocephalus versi- 
color that was distributed everywhere in the Gobi, and 
in many different macrohabitats used a rather limited 
range of microconditions (Fig. 2). In this respect its 
spatial niche was not broader than the niche of E. mul- 
tiocellata, and was even narrower in comparison with 
E. przewalskii. (Fig. 5). Eremias przewalskii which 
inhabits a rather limited range of macrohabitats is 
characterized by relatively broad requirements for 
microconditions along the first discriminant axis 
which represents the main direction of spatial segre- 
gation of lizard species. 

The structure of ecological space occupied by 
three lizard species has a complex character, explain- 
ing both macrohabitat and microhabitat segregation of 
spatial niches. Primarily, each discriminant axis char- 
acterizes the range of variation of microconditions in 
the study area. This can characterize macrohabitats 
only if variation in a set of variables included into 
analysis reflects the macrohabitat variation. In our 
case DF1 and DF2 possess such a feature: DF1 char- 
acterizes better the between-habitat component of 
spatial niche segregation and DF2 characterizes more 



the within-habitat component. Three species of lizards 
share microconditions in two-dimensional ecological 
space, so that the niche centers appeared to be almost 
equally distant from the geometrical center of the 
model (Fig. 5). The two species of Eremias lizards 
share ecological space to a greater extent than each 
Eremias with P. versicolor. Niche overlap between 
species was less than 15%. 

It seems difficult to speculate about processes that 
led to such spatial relationships. Interspecific compe- 
tition could contribute to spatial segregation of Ere- 
mias lizards in past, and could determine the low level 
of niche overlap. The competitive relationship 
between E. przewalskii and P. versicolor is unclear. In 
a pair of neighboring grid areas at one location within 
one habitat type when one species is in high density, 
the other one is usually at low density and vise-versa. 
However, there were no correlation between popula- 
tion densities of these two dominant species when all 
grid areas within one habitat type were put together. 
This result is easily explainable because different 
localities must have different upper limits of species 
densities according to the local environmental capaci- 
ties. However, the climatic conditions of Gobi Desert 
are so unfavored for reptiles (extremely low winter 
temperatures, great interannual, between- and within- 
seasonal contrasts in precipitation and temperatures) 
that it appears unrealistic to assume the space of 
resources is saturated by individuals, and that popula- 
tions are at equilibrium. 

Acknowledgments 

We are very grateful to Dr. David Ward (Ben-Gurion 
University of the Negev) for helping with the manu- 
script and to Dr. Natalia B. Ananjeva (Zoological 
Institute of St. Petersburg) for reading the manuscript. 
This is publication no. 12510-5241-9 of IEE and no. 
113ofRSC. 

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aspects of ecology of Mongolian desert lizards]. 
Nature and biological resources of Mongolian Peo- 
ple's Republic. Moscow. [Abstr.] (In Russian). 

Borkin, L. J., and D. V.Semenov. 1984. [Distribution 
of Phrynocephalus versicolor in South Mongolia]. 
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Brown J. H. 1984. On the relationship between abun- 
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Brown J. H. 1995. Macroecology. The University of 
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Case, T. J. 1983. Niche overlap and the assembly of 
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Fuentes, E. R. 1976. Ecological convergence of lizard 
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Henle, K. 1989. Ecological segregation in an assem- 
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Inger, R. F, and R. K. Colwell. 1977. Organization of 
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Kazantseva, T I. 1986. [Distribution and dynamics of 
the overground phytomass]. Pp. 106-114 In V.E. 
Sokolov and PD. Gunin (eds.). [Deserts of Trans- 
Altai Gobi]. Nauka, Moskva. (In Russian) 

Lavrenko, E. M. 1978. [On vegetation of steppes and 
deserts of Mongolian People's Republic]. Problemy 
osvoeniya pustyn. Ashkhabad 1:3-18. (In Russian). 

Liu Nai-Fa, Ren-De Li, and Xiao-Cheng Liang. 
1992. [Community structure of desert lizard in Gansu 
Province]. Acta Zoologica Sinica 38(4):377-384. (In 
China). 

Millado, J., F Amores, F F. Parreno, and F Hiraldo. 
1975. The structure of a Mediterranean lizard com- 
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Munkhbayar, Kh. 1976. [Amphibians and reptiles of 
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(In Mongolian). 

Murzaev, E. M. 1952. [Mongolian People Republic]. 
472pp. Geografgiz. Moskva. (In Russian) 

Pianka, E. R. 1973. The structure of lizard communi- 
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Pianka, E. R. 1975. Niche relations of desert lizards. 
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Podtyazhkin, O. I. and V. N. Orlov. 1986. [Faunoge- 
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regionalization of Mongolian People's Republic]. 



Institute of Animal Evolutionary Morphology and 
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Semenov, D. V. 1991. [Variations of characteristics 
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Semenov, D. V, and L. J. Borkin. 1986. [Amphibians 
and reptiles]. Pp. 1 14-119. In V E. Sokolov and P. D. 
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2001 



Asiatic Herpetological Research 



Vol. 9, pp. 122-129 



The Morphology and Size of Blood Cells of Lacerta rudis bithynica 
Murat Sevinc 1 and Ismail H. UGurta§ 

Uludagg University, Science and Art Faculty, Department of Biology, 16059 Bursa, Turkey. To whom 
correspondence should be addressed e-mail: smurat® uludag.edu.tr 

Abstract.- In this study, the morphology of the blood cells of Lacerta rudis bithynica is described using Wright's 
technique. The sizes of erythrocytes and their nuclei, leukocytes (monocytes, lymphocytes, basophils, 
neutrophils, and eosinophils), and thrombocytes of L. rudis bithynica were measured using an ocular micrometer 
at a magnification of 1600X with an oil immersion objective. The results of this study are compared with 
previous work on other reptile species. 



Key words.- 
morphology. 



Lacertidae, Lacerta rudis bithynica, erythrocyte, leukocyte, thrombocyte, measurement. 



Introduction 

The first studies on the blood of reptiles described the 
cellular structures, often comparing them with those 
of other vertebrates. Literature on the haematology of 
reptilian blood is based on few studies and is usually 
concerned with European species (Saint Girons, 
1970). 

Recent studies have concentrated on single species 
(Tiliqua sp., Cannon et al., 1988; Cyrtopodion sca- 
brum, Canfield and Shea, 1996). Various authors have 
described different circulating blood cells of different 
reptile species (Taylor and Kaplan, 1961; Heady and 
Rogers ,1962; Hartman and Lessler, 1964; Szarski 
and Czopek, 1966; Duguy, 1970; Saint Girons, 1970, 
Cannon et al., 1988, Canfield and Shea, 1996). Other 
authors have studied seasonal (Hutton, 1960; Cline 
and Waldman, 1962; Haggag and et. al, 1966) or sex- 
ual (Altland and Thompson, 1962) variation in the 
number of blood cells of different reptile species. In 
addition, authors have studied the number of blood 
cells of different reptile species (Charipper and Davis, 
1932; Baker and Cline, 1932; Hutton, 1961; Altland 
and Thompson, 1962; Hutchinson and Szarski, 1965; 
Engbretson and Hutchinson, 1976). Finally, authors 
have studied haemoglobin and hematocrit content of 
blood and hematopoiesis of different reptile species 
(Altland and Thompson, 1958; Hutton, 1961; Goin 
and Crawford, 1965; Engbretson and Hutchinson, 
1976; Newlin and Ballinger, 1976). 

In Turkey, haematological studies have been con- 
ducted on humans and economically important ani- 
mals. However, there are no haematological studies 
on the Turkish reptiles. 



In this study, our aim is to describe and measure 
blood cells (erythrocyte, leukocyte, thrombocyte) of 
Lacerta rudis bithynica (Squamata: Lacertidae). This 
study is the first of its kind on a Turkish species. 

Material and Methods 

In this study, 31 individuals (17 male) of Lacerta 
rudis bithynica (Squamata: Lacertidae) were exam- 
ined. The study was carried out between June and 
August 1998. The specimens were collected from 
Uludagg (Bursa) at an altitude of 1745 m. Blood was 
obtained by cardiac puncture of the lizards (Canfield 
and Shea, 1988). Immediately after the blood was 
obtained in heparinized capillary tubes, the blood 
smears were prepared. Three to five blood smears 
were prepared per individual. The smears were air- 
dried and stored until stained with Wright's stain 
(Hartman and Lessler, 1964). Twelve drops of 
Wright's stain were dropped on the slides and allowed 
to remain on the slide one and a half minutes before 
rinsing with a phosphate buffer (pH=6.5). The slides 
were allowed to stand for ten minutes at room temper- 
ature and were then washed with distilled water and 
allowed to dry. 

On each slide fifty mature erythrocytes and their 
nuclei, ten thrombocytes, and ten leukocytes (mono- 
cyte, lymphocyte, eosinophil, basophil and neutro- 
phil) were measured by means of an ocular 
micrometer at a magnification of 1600 x with an oil 
immersion objective. Cell sizes were calculated from 
the measurements. 



2001 



Asiatic Herpetological Research 



Vol. 9, 



123 




10/"" 



Figure 1 . Erythrocytes, an infected erythrocyte and a 
mitosis dvision. 

Results 

I. Erythrocytes 

Erythrocytes are nucleated, oval cells. Their nuclei are 
also oval, more or less regular and centrally located 
(Fig. 1). The cytoplasm of mature erythrocyte 
appeared light and dark pink and homogeneous under 
Wright's stain. The nuclei of mature erythrocytes are 
chromophilic. In some blood smears, immature eryth- 
rocytes are seen. They are characterised by a rounded 
form, blue cytoplasm and a large nucleus. Mitotic fig- 
ures are also present and in some smears, intracorpus- 
cular parasites are seen (Fig. 1). Parasites alter the 
shape and size of erythrocytes remarkably. When 
intracorpuscular parasites are seen, immature erythro- 
cytes and mitotic figures are abundant (Fig 1 ). Intrac- 
orpuscular parasites alter the shape and size of 
infected erythrocytes. The shape and size of other 
erythrocytes that are not infected by intracorpuscular 
parasites are normal. 



20 
15 



•s io 

s 





14 41 15 05 1 


5 71 






[6.17 [630 


M6M 



I Erythrocyte 
I Nucleus 



June 



July August 



Figure 2. Erythrocyte and nucleus lengths of Lacerta 
rudis bithynica qnq\ three months. 



to 
8 
6 
4 
2 




7.63 8.08 S 24 ■ 

BR ™ 


[3.5s [3 55 lj_ 74 



Erythrocyte 
Nucleus 



June 



July 



August 



Figure 3. Erythrocyte and nucleus widths of Lacerta 
rudis bithynica over three months. 

In June mean length of mature erythrocytes was 
14.41 urn (±0.77 standard deviations, with a range of 
12.20-16.47 urn). In July, the mean length of mature 
erythrocytes was 15.05 urn (±0.79, 12.81-17.08 pm). 
In August, the mean length of mature erythrocytes 
was 15.71 urn (±0.79, 12.81-18.30 um). Other mea- 
surements are given in Tables 1, 2 and 3. There are no 
significant differences in erythrocyte and nucleus 
sizes between females and males. Based on Tables 1 , 
2, and 3 and Figs. 2 and 3, it appears that there were 
little monthly variations in erythrocyte and nucleus 
sizes among June, July and August. 

II. Leukocytes 

1. Eosinophils. In blood smears stained by Wright 
technique, eosinophils are circular, and the cytoplasm 



Table 1. Erythrocyte dimensions of Lacerta rudis bithynica with the standard deviations in June. EL: Erythrocyte 
length; EW: Erythrocyte width; ES: Erythrocyte size;NL: Nucleus length; NW: Nucleus width; NS: Nucleus size. 



EL(um) 



EW (um) 



EL/EW 



ES (Mm 2 ) 



NS/ES 



Maximum 


16.47 ±0.77 


9.15 ±0.48 


2.27 ±0.13 


110.41 ±8.00 


0.30 ± 0.02 


Minimum 


12.20 ±0.77 


6.71 ± 0.48 


1.42 ±0.13 


64.26 ± 8.00 


0.13 ±0.02 


Mean 


14.41 ±0.77 


7.63 ± 0.48 


1.89 ±0.13 


86.46 ± 8.00 


0.20 ± 0.02 




NL(um) 


NW (um) 


NL/NW 


NS (Mm 2 ) 




Maximum 


7.32 ± 0.40 


4.88 ± 0.34 


2.20 ±0.19 


24.53 ±2.19 




Minimum 


4.88 ± 0.40 


3.05 ± 0.34 


1.25 ±0.19 


11.68 ±2.19 




Mean 


6.17 ±0.40 


3.55 ± 0.34 


1.75 ±0.19 


17.25 ±2.19 





Vol. 9, p. 124 



Asiatic Herpetological Research 



2001 



Table 2. Erythrocyte dimensions of Lacerta rudis bithynica\oqe\!r\er with the standard deviations in July. EL: Eryth- 
rocyte length; EW: Erythrocyte width; ES: Erythrocyte size; NL: Nucleus length; NW: Nucleus width; NS: Nucleus 



size. 



EL ( M m) 



EW (pm) 



EL/EW 



ES (nm 2 ) 



NS/ES 



Maximum 17.08 ±0.79 9.15 ±0.49 2.16 ±0.12 122.68 ±8.95 0.28 ± 0.02 

Minimum 12.81 ±0.79 6.71 ± 0.49 1.53 ±0.12 67.47 ± 8.95 0.13 ±0.02 

Mean 15.05 ±0.79 8.08 ± 0.49 1.86 ±0.12 95.62 ± 8.95 0.18 ±0.02 



NL(um) 



NW (Mm) 



NL/NW 



NS (Mm 2 ) 



Maximum 
Minimum 



Mean 



7.30 ± 0.70 4.27 ± 0.32 



5.40 ± 0.70 3.05 ± 0.32 



6.30 ± 0.70 



3.55 ± 0.32 



2.40 ±0.17 
1.33 ±0.17 



1.79 ±0.17 



24.53 ±2.38 
13.14 ±2.38 
17.67 ±2.38 



is stained light red. Eosinophils contain circular to 
elongate cytoplasmic granules stained brilliant red 
(Fig 4). Eosinophils are different from neutrophils in 
that, eosinophils' granules are stained bright red and 
neutrophils' granules were stained dim red. 

In June, the mean diameter of eosinophils was 
12.82 pm, (±1.71 standard deviations, with a range of 
9.93-15.25 um). In July, the mean diameter was 
13.29 um (±1.25, 10.98-15.25 um). In August, the 
mean diameter was 13.80 pm (±1.37, 12.20-15.25 
um). 

There were no significant differences in eosinophil 
diameters between females and males. Based on 
Tables 4, 5, and 6 and Fig. 8, it appears that there was 
little monthly variation in the diameter of eosinophils 
during the three months. 

2. Basophils. Basophils are easily recognised. They 
are small and circular cells. Nuclei stained blue by 
Wright technique are commonly obscured by chro- 
mophilic circular granules. These cytoplasmic gran- 
ules are large and stained dark purple. In the blood 



<p 

Erythrocytes j^k 




•lO/*"- 



smears, they resemble mulberries (Fig. 5). The gran- 
ules are so dense that nucleus stained dim blue is 
rarely seen. 

In June, the mean diameter of basophils was 8.55 
um, (±0.61 standard deviations, with a range of 7.32- 
9.15 pm). In July, the mean diameter was 9.02 pm 
(±0.24, 8.54-9.15 pm). In August, the mean diameter 
was 9.00 pm (±0.45, 7.93 -10.37 pm). 

There were no significant differences in basophil 
diameters between females and males. Based on 
Tables 4, 5, and 6 and Fig. 8, it appears that there was 
little monthly variation in the diameter of basophils 
during the three months. 

3. Neutrophils. Neutrophils are circular cells like 
eosinophils (Fig. 6). These cells are also called as het- 
erophils. They have cytoplasmic granules. The gran- 
ules are circular and stained dim red. Cytoplasm is 
stained light red. 

In June, the mean diameter of neutrophils was 
1 0. 1 5 pm, (± 1 . 1 9 standard deviations, with a range of 
9.15-13.42 pm). In July, the mean diameter was 10.49 




10 /u.n^ 



Figure 4. Erythrocytes and an eosinophil. 



Figure 5. Erythrocytes and a basophil. 



2001 



Asiatic Herpetological Research 



Vol. 9, p. 125 



Table 3. Erythrocyte dimensions of Lacerta rudis bithynica together with the standard deviations in August. EL: 
Erythrocyte length; EW: Erythrocyte width; ES: Erythrocyte size; NL: Nucleus length; NW: Nucleus width; NS; 
Nucleus size. 





EL (urn) 


EW (urn) 


EL/EW 


ES (urn 2 ) 


NS/ES 


Max 


18.30 ±0.76 


9.76 ± 0.48 


2.33 ±0.12 


140.20 ±8.96 


0.30 ± 0.02 


Min 


12.81 ±0.76 


6.71 ± 0.48 


1.57 ±0.12 


73.60 ± 8.96 


0.12 ±0.02 


Mean 


15.71 ±0.76 


8.24 ± 0.48 


1.91 ±0.12 


101.72 ±8.96 


0.18 ±0.02 




NL (urn) 


NW (pm) 


NL/NW 


NS (pm 2 ) 




Max 


7.93 ± 0.51 


4.27 ± 0.31 


2.40 ± 0.20 


24.53 ± 2.20 




Min 


4.88 ± 0.51 


3.05 ±0.31 


1.28 ±0.20 


11.68 ±2.20 




Mean 


6.48 ±0.51 


3.74 ±0.31 


1.74 ±0.20 


19.05 ±2.20 





Table 4. Leukocyte and thrombocyte measurements of L rudis bi/bynica with the standard deviations in June. 





Lym- 
phocyte 
(Mm) 


Monocyte 
(Mm) 


Neutrophil 
(Mm) 


Basophil 
(Mm) 


Eosinophil 
(Mm) 


Thrombo- 
cyte 
Length 
(Mm) 


Throm- 
bocyte 
Width 
(Mm) 


Max 


8.54 ± 0.81 


12.81 ± 


13.42 ± 


9.15± 


15.25 ± 


7.32 ± 0.49 


4.88 ± 






1.23 


1.19 


0.61 


1.71 




0.54 


Min 


4.27 ±0.81 


9.32 ± 1.23 


9.15 ± 1.19 


7.32 ± 
0.61 


9.93 ± 1.71 


4.88 ± 0.49 


3.05 ± 
0.54 


Mean 


6.12 ±0.81 


11.10± 


10,15 ± 


8.55 ± 


12.82 ± 


6.12 ±0.49 


3.72 ± 






1.23 


1.19 


0.61 


1.71 




0.54 



Table 5. Leukocyte and thrombocyte measurements of L. rudis biibynica w\tk the standard deviations in July. 





Lym- 
phocyte 
(Mm) 


Monocyte 
(Mm) 


Neutrophil 
(Mm) 


Basophil 
(Mm) 


Eosinophil 
(Mm) 


Thrombo- 
cyte 
Length 
(Mm) 


Throm- 
bocyte 
Width 
(Mm) 


Max 


7.32 ± 0.41 


15.25± 


12.20 ± 


9.15± 


15.25± 


7.32 ± 0.52 


4.88 ± 






1.5.0 


0.97 


0.24 


1.25 




0.42 


Min 


6.10 + 0.41 


9.15 ± 1.50 


8.54 ± 0.97 


8.54 ± 
0.24 


10.98 ± 
1.25 


6.10 ±0.52 


3.66 ± 
0.42 


Mean 


6.62 ±0.41 


11.46 ± 


10.49 ± 


9.02 ± 


13.29 ± 


6.62 ±0.52 


4.08 ± 






1.50 


0.97 


0.24 


1.25 




0.42 



Vol. 9, p. 126 



Asiatic Herpetological Research 



2001 



Table 6. Leukocyte and thrombocyte measurements of L rudis bithynicamVn the standard derivations in August. 





Lym- 
phocyte 
(Mm) 


Monocyte 
(urn) 


Neutrophil 
(pm) 


Basophil 
(pm) 


Eosinophil 
(pm) 


Thrombo- 
cyte 
Length 
(pm) 


Throm- 
bocyte 
Width 
(pm) 


Max 


9.15 ±0.58 


15.25± 


12.20± 


10.37 ± 


14.03 ± 


7.32 ± 0.35 


5.49 ± 






1.29 


0.97 


0.45 


1.37 




0.49 


Min 


6.10 ±0.58 


9.15± 1.29 


8.54 ± 0.97 


7.93 ± 0.45 


12.20 ± 
1.37 


5.49 ± 0.35 


3.05 ± 
0.49 


Mean 


6.53 ± 0.58 


11.21 ± 


10.77 ± 


9.00 ± 0.45 


13.80 ± 


6.33 ± 0.35 


4.22 ± 






1.29 


0.97 




1.37 




0.49 



um (±0.97, 8.54-12.20 pm). In August, the mean 
diameter was 10.77 um (±0.97, 8.54-12.20 um). 

There were no significant differences in neutrophil 
diameters between females and males. Based on 
Tables 4, 5, 6 and Fig.8, it appears that there was little 
monthly variation in diameter of neutrophils during 
the three months. 

4. Monocytes . Monocytes are round cells with round 
nuclei. The cytoplasm is stained blue and the nucleus 
is stained purple by Wright's technique. The mono- 
cyte's cytoplasm is more abundant than lymphocyte's 
cytoplasm. Nuclei vary in shape (Fig. 7). Nuclei may 
be nodular, but they are not lobular like granulocytes. 
Sometimes nuclei are horseshoe-shaped. 

In June, the mean diameter of monocytes was 
1 1.10 um, (±1.23 standard deviations, with a range of 
9.32-12.81 pm). In July, the mean diameter was 1 1.46 
pm (±1.50, 9.15-15.25 pm). In August, the mean 
diameter was 1 1 .2 1 um (± 1 .29, 9. 1 5- 1 5.25 pm). 

There were no significant differences in monocyte 
diameters between females and males. Tables 4, 5, 6 
and Fig. 8 show that there was little monthly variation 
in the diameter of monocytes during the three months. 



5. Lymphocytes . Lymphocytes are round cells like 
monocytes, but smaller (Fig. 7). The nuclei contain 
many parts of the cell. The nucleus is stained purple, 
cytoplasm, rarely seen, is stained blue. 

In June, the mean diameter of lymphocytes was 
6.12 um, (±0.81 standard deviations, with a range of 
4.27-8.54 um). In July, the mean diameter was 6.62 
pm (±0.41, 6.10 - 7.32 um). In August, the mean 
diameter was 6.53 um (±0.58, 6.10-9.15 pm). 

There were no significant differences in lympho- 
cyte diameters between females and males. Based on 
Tables 4, 5, 6 and Fig. 8, it appears that there was little 
monthly variation in diameter of lymphocytes during 
the three months. 

III. Thrombocytes 

Thrombocytes are small cells like lymphocytes, but 
they are oval and smaller than lymphocytes (Fig. 9). 
Their nuclei are highly chromophilic and stained pur- 
ple. Cytoplasm is rarely seen. 

In June, the mean length of thrombocytes was 6.12 
pm (±0.49 standard deviations, with a range of 4.88- 
7.32 pm). In July, the mean length of thrombocytes 






~^~ 




Jfc 




¥ ^SfeP^ Neutrophil 



Erythrocytes 



, * f 



\0f*-* 



Lymphocyte 




lO/*"" 



Figure 6. Erythrocytes and a neutrophil. 



Figure 7. Erythrocytes, a monocyte and a lymphocyte. 



2001 



Asiatic Herpetological Research 



Vol. 9. p. 127 




□ June 
■ July 
E3 August 



lymphocyte monocyte neutrohpil basophil eosinophil 

Leukocytes 

Figure 8. Leukocyte diameters of Lacerta rudis bithynica over three months. 



♦ 










Thrombocytes 



Ery throe ytes 



\Of*-r 



\ 



Figure 9. Erythrocytes and thrombocytes. 



8 

6 12 

ml- 




662 

1408 


6.33 


3.72 


I 4 - 




■ 



I Length 
I Width 



June 



August 



Figure 10. Thrombocyte sizes of Lacerta rudis 
bithynica over three months. 

was 6.62 um (±0.52, 6.10-7.32 urn). In August, the 
mean length of thrombocytes was 6.33 um (0.35, 
5.49-7.32 um). 

In June, the mean width of thrombocytes was 3.72 
(±0.54 standard deviations, with a range of 3.05-4.88 



um). In July, the mean width of thrombocytes was 
4.08 um (±0.42, 3.66-4.88 um). In August, the mean 
width of thrombocytes was 4.22 um (0.49, 3.05-5.59 
Mm). 

There were no significant differences in thromb- 
ocytes sizes between females and males. Based on 
Tables 4, 5, 6 and Fig. 10, it appears that there was lit- 
tle monthly variation in thrombocytes sizes during the 
three months. 

Discussion 

Investigations carried out by various authors (Hart- 
man and Lessler, 1964; Szarski and Czopek, 1966; 
Saint Girons, 1970) reported that the sizes of the 
erythrocytes vary in members of the four orders of 
reptiles. Within the class Reptilia, the largest erythro- 
cytes are seen in Sphenedon punctatus, turtles and 
crocodilians. The erythrocytes of lizards vary greatly 
in size depending on the family and sometimes even 
within one family (Saint Girons, 1970). The smallest 
erythrocytes are found in the lizard family Lacertidae 
(Saint Girons, 1970). 

In the present study, erythrocyte morphology and 
the results of erythrocytes sizes (Table 1, 2 and 3) are 
agreement with the other results carried out by Hart- 
man and Lessler (1964), Szarski and Czopek (1966), 
and Saint Girons (1970). 

In one of the studies on the leukocytes of the 
rough tail Gecko Cyrtopodion scabrum, a bright-field 
and phase-contrast study Cannon et al. (1996), 



Vol. 9, p. 128 



Asiatic Herpetological Research 



2001 



reported that the neutrophils were not observed, but 
the other leukocytes were observed. 

Another study on morphological observations on 
the erythrocytes, leukocytes and thrombocytes of blue 
tongue lizards by Canfield and Shea (1988) reported 
that all types of leukocytes were observed. 

Saint Girons (1970) and Canfield and Shea (1988) 
divided granulocytes into neutrophils, basophils and 
eosinophils on the basis of light microscopy. However 
Cannon et al. (1996) divided granulocytes into baso- 
phils and eosinophils on the basis of bright-field and 
phase-contrast microscopy. 

Heady and Rogers (1962) divided leukocytes into 
neutrophils, small acidophils, eosinophils, lympho- 
cytes and monocytes on the basis of light microscopy 
in Pseudemys elegans. They, except for monocytes, 
gave the sizes of leukocytes and reported that eosino- 
phils and neutrophils were numerous than the other 
leukocytes. Taylor and Kaplan (1961) also divided 
leukocytes into neutrophils, basophils, eosinophils, 
lymphocytes and monocytes on the basis of light 
microscopy in turtles. 

In this study, it appears that on the basis of light 
microscopic findings there are three main types of 
granulocytes and two types of agranulocytes in L. 
rudis bithynica and also the size of all kinds of leuko- 
cytes are given in Table 4, 5 and 6. 

In the present study, the descriptions and sizes 
(Table 4, 5 and 6) of thrombocytes are comparable to 
other descriptions. Canfields and Shea (1988) 
reported that thrombocyte morphology at the light 
microscopic level is influenced by the degree of 
aggregation and degranulation. Saint Girons (1970) 
reported that thrombocytes are small, oval cells char- 
acterised by elongate, centrally located highly chro- 
mophilic nuclei. The cytoplasm is almost colourless 
(faintly acidophilic) and hence difficult to see in a 
blood smears. Taylor and Kaplan (1961) reported the 
same findings in turtles. 

Literature Cited 

Altland, P. D. and E. C. Thompson. 1958. Some fac- 
tors affecting blood formation in turtles. Proceedings 
for the Society of Experimental Biology and Medi- 
cine. 99:456-459. 

Baker, E. G. S. and L. E. Kline. 1932. Comparative 
erythrocyte count of representative vertebrates. Pro- 
ceedings of the Indiana Academy of Sciences 41:417- 
418. 

Canfield, P. J. and G. M. Shea. 1988. Morphological 
observations on the erythrocytes, leukocytes and 



thrombocytes of blue tongue lizards (Lacertilia: Scin- 
cidae, Tiliqua). Anatomia, Histologia, Embryologia. 

17:328-342. 

Cannon, M. S., D. A. Freed and P. S. Freed. 1996. The 
leukocytes of the roughtail gecko Cyrtopodion sca- 
brum: a bright-field and phase-contrast study. Anato- 
mia, Histologia, Embryologia. 25:1 1-14. 

Charipper, H. A. and D. Davis. 1932. Studies on the 
arneth count. A study of the blood cells of Pseudemys 
elegans with special reference to the polymorphonu- 
clear leukocytes. Q. J. Experimental Physiology. 
21:371-382. 

Cline, M. J. and T. A. Waldmann. 1962. Effect of tem- 
perature on red cells in the alligator. Proceedings for 
the Society of Experimental Biology and Medicine. 
111:716-718. 

Duguy, R. 1970. Numbers of blood cells and their 
variations. Pp. 93-104. In Gans (ed.), Biology of the 
Reptilia, Vol. 3, Morphology C. Academic Press, New 
York. 

Engbretson. G. A. and V H. Hutchinson. 1976. Eryth- 
rocyte count, hematocrit and haemoglobin content in 
the lizard Liolaemus multiformis. Copeia 1976:186. 

Goin, C. J. and C. G. Jackson. 1965. Haemoglobin 
values of some amphibians and reptiles from Florida. 
Herpetologica 21:1 45-146. 

Haggag, G., and et al. 1966. Hibernation in reptiles II. 
Changes in blood glucose, haemoglobin, red blood 
cells count, protein and nonprotein nitrogen. Compar- 
ative Biochemistry and Physiology. 17:335-339. 

Hartman, F. A., and M. A. Lessler. 1964. Erythrocyte 
measurements in fishes, amphibians and reptiles. Bio- 
logical Bulletin. 126:83-88. 

Heady, J. M. and T. E. Rogers. 1963. Turtle blood cell 
morphology. Proceedings of the Iowa Academy of 
Sciences. 69:587-590. 

Hutchinson, V H. and H. Szarski. 1965. Number of 
erythrocytes in some amphibians and reptiles. Copeia 
1965:373-375. 

Hutton, K. E. 1960. Seasonal physiological changes 
in the red eared turtle Pseudemys scripta elegans. 
Copeia 1960:360-362. 

Hutton, K. E. 1961. Blood volume, corpuscular con- 
stants and shell weight in turtles. American Journal of 
Physiology 200:1004-1006. 

Newlin, M. E. and R. E. Ballinger. 1976. Blood hae- 
moglobin concentration in four species of lizards. 
Copeia 1976:392-394. 

Saint Girons, M. C. 1970. Morphology of the circu- 
lating blood cells. Pp. 73-91. In Gans (ed.), Biology of 



2001 Asiatic Herpetological Research Vol. 9, p. 1 29 



the Reptilia, Vol. 3, Morphology C. Academic Press, 
New York. 

Szarski, H. and G. Czopek. 1966. Erythrocyte diame- 
ter in some amphibians and reptiles. Bulletin de 
l'Academie Polonaise des Science. Classe 2. Serie des 
Sciences Biologiques. 14(6):437-443. 

Taylor, K. and H. M. Kaplan. 1961. Light microscopy 
of the blood cells of Pseudemyd Turtles. Herpetolog- 
ica 17:186-192. 



2000 



Asiatic Herpetological Research 



Vol. 9, pp. 130-138 



Buccopharyngeal Morphology and Feeding Ecology of Microhyla ornata 

Tadpoles 

Muhammad S. Khan 

7740 NW 47th St., Lauderhill, FL 33351 USA 

Abstract.- The oropharyngeal morphology of Microhyla ornata tadpoles is described. Anatomical peculiarities 
are correlated to feeding ecology. Microhylid anatomical features are discussed and compared with ranoid 
tadpoles. 

Key words.- Amphibia, Salientia, Microhylidae, Microhyla ornata, larval oropharyngeal morphology, feeding 
ecology. 



Introduction 

The common southeast Asian narrow-mouth frog 
Microhyla ornata is widely distributed throughout 
Punjab, Sindh. N.W.F.P. and Azad Kashmir (Khan, 
1974; 1979; 1988; Khan andTasnin, 1987). Normally, 
its small size makes it inconspicuous and difficult to 
detect in the field. However, after a summer heavy 
downpour, the characteristic rasping call of Microhyla 
ornata is easily distinguishable from rest of the local 
amphibians (Khan and Malik, 1987b). Microhyla 
ornata readily takes refuge under vegetation, leaf lit- 
ter, logs, stones, in holes and fissures in the ground, 
and often is mistaken as a juvenile of some larger spe- 
cies. 

At mid-monsoon, M. ornata invades large water 
bodies, which, by this time, are filled with water and 
have developed thick planktonic growth. Solitary 
males perch well away from water among marginal 
vegetation to call. Eggs are laid in patches of jelly, 
which float at water surface as "egg-rafts" (Khan, 
1982b). 

The present study describes oropharyngeal morphol- 
ogy of Microhyla ornata tadpole, at Stage 35 and cor- 
relate it to the tadpole's feeding ecology. Moreover, it 
is compared with already known morphologies of 
sympatxic tadpoles belonging to the genera Bufo and 
Rana (Khan and Malik, 1987a; Khan and Mufti, 
1994b, 1995). 

Material and Methods 

Tadpoles for the present study were collected during 
the summers of 1986-88 from different localities 
along the northwestern border of Rabwah City (Khan 
and Malik, 1987b). They were netted at midstream 
with a hand net. Microhyla ornata tadpoles from 
Ghakkhar, District Gujranwala, Punjab, Pakistan 



(Khan, 1974) and Azad Kashmir (Khan, 1979), were 
used as comparative material. 

Collection, preservation and storage methodology 
followed Khan (1982b), while surgical procedures 
and descriptive terminology are from Khan and Malik 
(1987a) and Khan and Mufti (1994b, 1995), except 
that for M. ornata tadpole, the lateral cuts through 
buccopharyngeal walls, must pass through mid-eye, 
unlike ranoid tadpoles where cuts pass below the eye. 
Fine particulate mucilage-trapped material accumu- 
lating in the filter cavities is cleared by a jet of water 
from an ordinary eyedropper. Drawings of the buc- 
copharyngeal surfaces were made with the help of 
camera lucida. 

Tadpoles at Stage 35 were selected, since at this 
stage, they have already attained maximum size and 
their characteristic organs are fully developed and 
functional. The tadpole at this stage is voraciously 
feeding and its digestive system is functioning at its 
full capacity. Shortly after this stage, metamorphic 
changes start occurring. 

For identification of Stage 35, tadpoles were com- 
pared with Khan's (1965) table of normal develop- 
ment. Data for present study are recorded from 10 
specimens. 

Description 

External morphology 

The tadpole's body is perfectly streamlined; the head 
is dorsoventrally depressed while its belly is laterally 
compressed and oval in dorsal profile. The snout is 
countersunk, displacing mouth anterodorsally. The 
tail is more than twice the length of the head and 
body. Broad caudal fins narrow abruptly in the poste- 






2000 



Asiatic Herpetological Research 



Vol. 9, p. 131 



buccal floor arena 
(BFA) 



preglottal 

papilla 

laryngeal disc 



ventral 
vellum 



filter cavity 




nfralabial papillae 
tongue analage 



buccal pocket 

lottis 

BFA papillae 



buccal pocket 
canal 



trachea 



esophagus 



Morphology of surgically exposed buccopharyngeal floor of Microhyla omata\adpo\e at Stage 35 (scale 



Figure 1. 
1 mm). 

nor half of the tail, passing into a delicate long flagel- 
lum. 

The body is widest at the level of laterally dis- 
posed small eyes. The nostrils are imperforate; their 
position is marked by heavily pigmented anterolateral 
pits lying just anterior to eyes. The mid- ventral spira- 
cle is close to posterior ventral end of the abdomen, 
with a distinct prespiracular valve (Khan 1982b). 

The horizontal mouth has a median U-shaped cleft in 
the middle of the lower lip which remains perma- 
nently open (Khan and Mufti, 1994a, Fig. 2). Pres- 
ence of iridiocytes in the abdominal wall, give it a 
characteristic silvery shine, which is lost within two to 
three weeks, on preservation. A median dorsal band of 
melanophores covers the brain and extends onto the 
base of eyes and the nasal pits. 

Measurements (in mm). Body length 5.2-5.8; tail 
length (including flagellum) 12.6-13.7; total length 
17.8-18.9; greatest breadth of body (at the level of 
eyes) 2.7-2.9; greatest depth of body (at level of spira- 



cle) 3.2-3.4; interorbital space 2.3-2.4; internarial 
space 0.5-1.95; tail muscle height (at base) 1.7-2.0; 
tail fin height (at midtail) 4-4.4; length of tail flagel- 
lum 2.0-2.3. 

Internal Morphology 

Buccal region. Khan and Mufti (1994b; 1995) distin- 
guished a tadpole's buccal cavity in two functional 
units: anterior food gleaning part and posterior food 
retrieving part. The dorsoventrally depressed head and 
peculiar position of the mouth in M. ornata tadpoles 
have affected the form and shape of the tadpole's buc- 
cal cavity; reducing the food gleaning part and widen- 
ing the food retrieval part which occupies most of the 
buccal region. 

Ventral buccal (Fig.l). The floor of the food glean- 
ing part consists of vertical U-shaped prelingual 
chamber which opens out through mouth at snout top 
and is lined by a series of three simple infralabial 
papillae. Posteriorly, it opens in food retrieval part of 



Vol. 9, p. 132 



Asiatic Herpetological Research 



2000 



postnarial ridge 



prenarial papilla 



nans 



buccal roof arena 
(BRA) 

postero- 
lateral 
BRA ridge 



narial papilla 
BRA papilla 



pressure 
cushions 




1 mm 



buccal 
roof 

glandular 
area 

I — dorsal 
vellum 



esophagus 



Figure 2. Morphology of surgically exposed buccopharyngeal floor o\Microhyla oma/a\adpo\e at Stage 35 (scale 
1 mm). 



the buccal. A non-papillated tongue analage, a conical 
thickening which is broader anteriorly pointed poste- 
riorly, guards the opening of the food gleaning part 
into the food retrieval part. The spacious food retrieval 
part forms the main buccal cavity. The buccal floor 
arena (BFA) is rectangular, laterally raised with a 
median shallow passage. A group of 12-14 fine 
tipped, large, flat BFA papillae lie on lateral sides of 
anterior end of trachea. A pair of smaller midpocket 
papillae lie at the level of mid-pockets. A buccal 
pocket is a long, narrow, club shaped longitudinal slit 
running anteroposteriorly on each lateral side of the 
BFA with a posterior narrow canal connecting it with 
the pressure cushions. Anterior, wider, parts of buccal 
pockets have 3-6 pre- and 5-8 post pockets, small, 
conical papillae. 

The trachea is a long cylindrical pipe that extends 
medially to the base of the BFA, carrying the glottis 
far anterior to the ventral velum and dividing it into 
lateral halves. The thin lipped glottis is 100% 
exposed, lies on a bulbous laryngeal disc, and is 



guarded by a long preglottal papilla which is tipped 
left. 

The broad ventral velum has a strong spicular sup- 
port. It covers about l/2-l/4th of the underlying bran- 
chial baskets and consists of three distinct long and 
deep filter cavities. The free margin of the velum is 
smooth, with a single broad projection above third fil- 
ter plate and is covered by a narrow strip of minute 
secretory pits. Rows of melanophores run along the 
lateral sides of the buccal arena and are aggregated on 
the sides of the tongue analage; a row runs along lat- 
eral sides of trachea. 

Dorsal buccal (Fig. 2). The food gleaning part of the 
buccal is roofed by a broad prenarial arena, which has 
a median V-shaped depression with an anterior pre- 
narial papilla and several pustules. The position of the 
imperforate naris is marked by a shallow depression 
from which a flat ribbon like twisted narial papilla 
hangs down in the buccal cavity. A thin delicate, nar- 
row, papillated, postnarial ridge dorsally delimits nar- 
ial region and the food gleaning part of the buccal. 



2000 



Asiatic Herpetological Research 



Vol. 9, p. 133 



laryngeal disc 



secretory ridg 



glottis 
trachea 




Figure 3. Underside of ventral velum showing details of filter cavities (diagramatic). 



The food retrieval part is roofed by a spacious, 
broader than long, buccal roof arena (BRA), which is 
featured mainly by a distinct posterolateral BRA 
ridge, the free margin of which is cut into flat fine 
tipped BRA papillae, increasing in length from out 
smallest, to inside longest, with blunts on their inner 
sides, while smaller are without blunts. The BRA sur- 
face is pustular. 

The broad glandular area of the buccal roof is 
divided into lateral rectangular halves and is covered 
with dense minute secretory pits. The dorsal velum 
narrows gradually mesoid, at mid-BRA it staggers to 
continue with that of other side. 

Branchial region. Branchial baskets are more than 
twice the length of the buccal. Three filter cavities are 
distinct in each branchial basket. The first filter cavity 
is largest while the third is smallest. The filter cavities 
are always packed with fllocular matter. The third is 
tilted outwards, partially blanketing the second. About 
half to one third of the filter cavities are covered with 
velum. A distinctly ridged oval torus (Fig. 3:1, II, III) is 
present in each filter cavity. The subvelar surface is 
profusely ridged with fine transverse secretory ridges 



(Fig. 3) that run in line with toric ridges, which are 
edged with fine secretory pores. 

A tight filter ruffle cover the surface of filter 
plates. The number of filter rows on filter plates vary 
from minimum 9 on the 4th ceratobranchial to 23 on 
second (Table 1 ). The filter ruffle is 3° dense with ter- 
tiary foldings. Successive filter rows abut across fully 
canopied deep filter canals. The filter ruffle covers 
both sides of second and third filter plates (Fig. 3: cb2, 
cb3). 

Three pressure cushions are distinct on posterolat- 
eral sides of the dorsal pharynx (Fig. 4). The first and 
second are four times longer than broad. The first, and 
outer-most, is continuous anteriorly with the buccal 
pocket of its side through a buccal pocket canal. 
Meanwhile, the third, innermost, is broadest and has a 
median hook-like appendage. Deep ciliary groove 
runs along posterior border of the pressure cushions 
towards esophageal orifice along posterior sides of the 
pressure cushions. 

The lungs at Stage 35 are well developed, each 
running along the dorsolateral sides of the abdominal 
cavity, extending to the posterior end of the abdomen. 
Anterior half of the lung is broad, with well-devel- 



Table 1. Branchial elements of Microhyla ornata tadpoles at Stage 35 (Ant=anterior aspect; B=breadth; cb=cerato- 
branchial; L=length; Post=posterior aspect). Data from 10 specimens, all measurements in mm. 



Ceratobranchial 



Filter plate 



Filter rows/side 



cb. 1 
cb.2 
cb.3 
cb.4 



3.5 



1.8 



3.7 


1.5 


2.5 


1.4 


1.9 


1.2 



Anterior 


Posterior 





15-18 


13-16 


20-23 


16-17 


13 


8-9 






Vol. 9, p. 134 



Asiatic Herpetological Research 



2000 



Table 2. Comparison of microhylid and ranoid tadpole. 



Character 



Microhylid 



Ranoid 



Head 

Belly 

Color 

Tail length 

Tail muscle 

Tail fins 

Tail tip 

Spiracle 

Belly wall 

Mouth 

Oral disc 

Nostrils 

Narial flap/papillae 

Infralabial papillae 

Lingual papillae 

Postnarial papillae 

Lateral ridge papilla 

Median ridge 

Buccal musculature 

Branchial basket 

Pharyngeal/buccal ratio 

Prenarial ridge 

BRA/BFA papillae 

BRA/BFA papillae 

Glottis 

Glottal disc 

Glottal position 

Glottal papilla 

Trachea 

Secretory tissue 

Subvelar secretory tissue 

Filter cavities 

Maximum filter rows 

Position of tori 

Filter cavity depth 



depressed 

compressed 

transparent 

2-3 times body 

narrow 

broad 

produced in a flagellum 

median ventral 

silver-shiny 

antero-dorsal 

absent 

imperforate 

flap 

simple 

absent 

small on a membrane 

absent 

absent 

poor 

longer than broad 

75% 

absent 

flat fine tipped 

form membranes 

1 00% exposed 

bulbous 

anterior to vellum 

present 

long 

single celled 

ridged 

3 

15-23 

all filter cavities 

as long as deep 



depressed 

depressed 

Drab + pattern 

2-2.5 times 

broad 

moderate 

round/pointed 

dextral 

transparent or drab 

anterior/antero-ventral 

present 

perforate 

papillae 

palmate 

present 

long solitary 

present 

present 

well developed 

broader than long 

45-50 % 

pustules or ridge 

short forked blunted 

distinct no membranes 

not or partiall exposed 

concealed 

posterior to vellum 

absent 

absent 

multicellular 

ridged/pitted 

2/3 

10-14 

absent or first cavity 

longer than deep 



2000 



Asiatic Herpetological Research 



Vol. 9, p. 135 



buccal pocket canal 



pressure cushions 




pressure cushion 
hook 



ciliary 
groove 



Figure 4. Morphologyofposterolateralpartofdorsal buc- 
copharyngeal region, showing details. 

oped air sacs, it gradually narrows down caudally and 
its terminal part is a dark pigmented cord. 

Ecological Correlates 

The adult Microhyla ornata is sharply contrasted from 
its sympatric ranoids in its choice of breeding site and 
mode of egg deposition. Usually its breeding sites are 
deep ponds, which develop rich planktonic bloom 
during summer. Such sites are quite restricted and rare 
in temperate Punjab, unlike sympatric ranoids who 
breed in temporal sites that dry in the summer. Inter- 
mittent unpredictable rains or irrigation seepage saves 
such populations of tadpoles (Khan and Malik, 
1987b). 

Table 1 and 2 summarize peculiar features of M. 
ornata tadpole necessary to suit its lentic habits: per- 
fectly streamlined transparent body and long broad 
finned tail with independently vibratile terminal fla- 
gellum, lateral eyes, median ventral spiracle, abdomi- 
nal silver-shine, are adaptations to nektic habits of the 
M. ornata tadpole. A school of tadpoles swimming at 
midstream is almost invisible from above the water 
due to tadpole's transparent bodies. Meanwhile 
abdominal coloration makes them invisible in water 
since the water surface viewed from inside appears 
shiny due to reflection of light from water. A vibrating 



tail flagellum and the jet of water from median ventral 
spiracle supports the microhylid tadpole at mid- 
stream, while its buoyancy is maintained by dorsally 
placed air filled lungs. A long broad finned tail helps 
the tadpole to react quickly to any stimulus including 
enemies and food in the water column above and 
below. 

Microphagous tadpoles are characterized by an 
exaggerated oropharyngeal region (Wassersug, 1980). 
All oropharyngeal modifications distinguishing 
microhylid tadpoles from bufonids and ranids are 
adaptations to microphagy: reduction of infralabial 
cartilage, corresponding reduction in prelingual and 
prenarial arenas: displacement of mouth to snout top; 
simple infralabial papillae; absence of lingual papil- 
lae; membranous postnarial and BRA papillary 
ridges; divided fine pitted buccal glandular zone; fine 
ridged subvelar and toric glandular surfaces; broad 
medially divided spicularly supported ventral velum; 
compact thick filter ruffle on broad and deep filter 
plates; deep filter cavities with distinct tori. These 
microhylid structures are part of obligate micropha- 
gus filter feeding technology. Similarly long tubular 
trachea, bulbous laryngeal cartilage, distinct anteri- 
orly displaced glottis with a preglottal papilla, and 
dorsal, long, air filled lungs are not only efficient parts 
of an efficient pulmonary aeration system, but at the 
same time, serve to maintain hydrostatic balance of 
the tadpole at mid-stream position. The complex mor- 
phology of pressure cushions and their connection 
with buccal pockets are a part of a system which 
maintains a sustained drainage of excessive buccal 
water in pressure cushions, providing necessary tur- 
gidity during each stroke of bucco-pharyngeal pump- 
ing. Turgid cushions act as pharyngeal valves in the 
process and play key role in sustained pumping of 
food-laden water current in buccopharyngeal passage. 

Morphologically, the Microhyla ornata tadpole is 
a perfect model suited for sustained midstream swim- 
ming. It has all the necessary oropharyngeal technol- 
ogy to filter and feed on the planktonic bloom in 
deeper ponds in temperate Punjab. 

Microhylid Tadpole Characters 

The microhylid tadpole is ranked as Type 2 in Orton's 
(1953) classification. It markedly differs in morpho- 
logically from ranoid Type 4 macrophagus tadpole 
(Table 2). Microhylid is the only tadpole, in Punjab 
riparian ecosystem, occupying unique midstream 
feeding niche, while rest of the sympatric ranoid tad- 
poles are bottom grazers and in no way compete with 
microhylid tadpole. Distinguishing morphological 
characteristics of Microhyla ornata tadpoles are: 



Vol. 9, p. 136 



Asiatic Herpetological Research 



2000 



External morphology 

1. Transparent body and tail, silver-shine on abdomen. 

2. Head dorso-ventrally depressed, body and tail later- 
ally compressed. 

3. Antero-dorsal mouth, without keratinized oral disc 
and other associated organs. 

4. Eyes prominent, laterally disposed on head. 

5. Tail broad finned, its tip produced into a terminal 
vibratile flagellum. 

6. Spiracle median-ventral, mid-abdominal, squarish 
opening, with a prespiracular flap. 

7. Anal tube straight, median-ventral anal opening. 

8. Schools of tadpoles swim at midstream schools, 
never rest at bottom. Capable of making spontaneous 
movements from midstream to darker parts of the 
pond to avoid intruders, shortly reappearing at the 
same midstream site. 

Internal morphology 

9. Opercular chamber extending to vent. 

10. Smooth broad ventral vellum, divided into right 
and left halves. 

1 1 . Long pipe like trachea carries, bulbous glottal car- 
tilage, 

for forward in buccal cavity. Glottis thin lipped, 100% 
exposed. 

12. Preglottal papillae guarding glottis. 

13. No lingual papillae. 

14. No lateral ridge papilla and median ridge. 

15. Pharyngeal region exaggerated, about 75 % of 
oropharyngeal region, branchial baskets large with 
distinct, deep filter cavi ties, which are usually full of 
fllocular matter in dissected tadpoles. 

16. Tight filter mesh, maximum number of filter rows 
13-23 on either sides of the ceratobranchials. 

17. Branchial food traps with microscopic openings of 
secretory glandular tissue borne on fine parallel ridges 
forming distinct crescentric torus in each branchial 
cavity. 

18. Imperforate nares with a foliaceous broad narial 
palp, descending into the buccal cavity. 

19. Ceratohyal with a ventrally directed lateral arm, 
and an an tero-posterior median arm. 

20. Broad based BRA and BFA papillae, which usu- 
ally coalesced to form fine membranes. 

21. Narrow lateral buccal pockets with distinct con- 
nection with pressure cushions, forming an elaborate 
system to control function of pressure cushions. 



Microhylids are phylogenatically connected with 
ranoids through tadpoles with intermediate morpholo- 
gies like Psedohemisus granulosa (Wassersug, 1984) 
and Otophryne robusta (Wassersug and Pyburn, 
1987). 

Discussion 

Independent vibratile distal caudal flagella are charac- 
teristic of mid-stream swimming microphagus tad- 
poles (Wassersug, 1980, 1989; Wassersug and Sperry, 
1977; Nishikawa and Wassersug, 1988, 1989; Hoff 
and Wassersug, 1986). Maintenance of midstream 
position is made possible by independent movements 
of the caudal flagellum and ventrally directed continu- 
ous water from the spiracle giving a sustained upward 
thrust (Khan, 1982a, 1991). Apart from morphologi- 
cal differences in structure of notochord and arrange- 
ment of caudal nerves, microhylids and ranoids differ 
in the site of generation of propulsive locomotory 
waves. In ranoids, waves are generated at the tail; the 
tip acts as a steer. In microhylids, waves are generated 
at the end of the caudal flagellum. 

The limited tail musculature and bulky form of the 
amphibian tadpole restricts its movements so that it 
cannot evade its potential enemies, fishes, niads, etc., 
(Khan and Mufti, 1994b; 1995). Amphibian tadpoles 
rely on a reduced conspicuousness. Its drab spotted 
pattern blends well against natural aquatic back- 
ground with moderate to thick vegetation (Caldwell, 
et al., 1981; Gatten et al., 1984; Kehr and Basso, 
1990; Khan and Mufti, 1994 b. 1995). The microhylid 
tadpole, which is exposed at midstream, solves this 
problem differently. Transparency of its body reduces 
its shadow at pond bottom and its abdominal shine 
blends well against water surface reflecting sun rays, 
making it invisible from inside pond as well from out 
side, to its predators. 

Microhyla omata has a larger buccal volume than 
ranoids. It constantly pumps large amounts of water to 
get food (Seale and Wassersug, 1979; Wassersug, 
1980). Due to its specialized feeding habits, several 
elements universally present in ranoid tadpoles are 
missing in its oropharyngeal morphology: median 
ridge, lateral ridge papillae and lingual papillae. 
Moreover, papillae in the food retrieval part are a part 
of particulate food guiding membrane rather particu- 
late food retrieving sieves. Moreover, microhylids 
have fine-ridged oral and branchial glandular system 
with fine pits, a specialization to entrap finest particu- 
late food. Deep filter cavities, long filter plates and 
tight filter mesh are more a part of food retrieval sys- 
tem rather respiratory in function (Wassersug and 
Murphy, 1987). Particulate food filtering capacity of 



2000 



Asiatic Herpetological Research 



Vol. 9, p. 137 



filtering system is enhanced by development of a fine- 
ridged torus in each filter cavity. A well-developed 
pulmonary system with long tubular trachea, a totally 
exposed guarded glottism, and inflated dorsal func- 
tional lungs are adaptations to the midstream sus- 
tained swimming and are efficient respiratory organs 
(Khan, 1991). 

Tadpoles of the microhylid genera Otophryne and 
Psdudohemisus show both ranoid as well as micro- 
hylid characteristics. Frogs of Family Microhylidae 
and Ranidae have distinctive adult morphology, how- 
ever definition of microhylid larva stands only on 
imperforate naris, since the ranoids have always per- 
forated naris (Wassersug, 1989). 

Tadpoles of M. ornata from Pakistan differ in mor- 
phological details from those collected from Thailand 
by Inger (1985) in having longer infralabial papillae, 
more filter rows, presence of postnarial membranous 
ridge, BRA and BFA membranous papillae, mesially 
divided dorsal buccal glandular zone, and a single 
preglottal papilla. Moreover, no silver shine is 
reported on the belly of tadpoles from Thailand, as is 
reported from India (Rao, 1917; Azad Kashmir 
(Khan, 1979) and District Jhang, Punjab Pakistan 
(Khan 1982a). Flower (1899) reported morphological 
differences among adults and larvae of this species 
from Malay peninsula and Siam, while Liu (1950) has 
reported dextral anal tube in tadpoles from China, 
which is straight in Pakistani tadpoles. These morpho- 
logical differences may refer to geographical races 
within this widely distributed southeast Asian species 
of narrow-mouth frogs. 

Literature Cited 

Caldwell, J. P., H. Thorp, and T. O. Jervey. 1981. 
Predator prey relationships among larval dragonflies, 
salamanders and anurans. Oecologia 46:285-289. 

Flower, S. S. 1899. Notes on a second collection of 
batrachians made in the Malay peninsula and Siam, 
from November 1896 to September 1898, with a list 
of the species recorded from these countries. Proceed- 
ings of the Zoological Society of London 1899:885- 
916. 

Gatten, R. E. Jr., H. P. Caldwell, and M. E. Stockard. 
1984. Anaerobic metabolism during intense swim- 
ming by anuran larvae. Herpetologica40:164-169. 
Hoff, K. S., and R. J. Wassersug. 1986. The kinemat- 
ics of swimming in larvae of the clawed frog, Xeopits 
laevis . Journal of Experimental Biology 122:1-12. 
Inger, R. F. 1985. Tadpoles of the forested regions of 
Borneo. Fieldiana Zoology, n.s. 1365(26): 1-89. 



Kehr, I, K„ and N. G. Basso. 1990. Description of the 
tadpole of Lysapsus limellus (Anura: Pseudidae) and 
some consideration on its biology. Copeia 1990:573- 

575. 

Khan, M. S. 1965. A normal table of Bufo stomaticus 
(Bufo melanostictus). Biologia (Lahore), 11:1 -39. 

Khan, M. S. 1974. Discovery of Microhyla ornata 
Dumeril and Bibron from Punjab, Pakistan. Biologia 
(Lahore) 20: 179- 180. 

Khan. M. S. 1979. On a collection of amphibians 
from northern Punjab and Azad Kashmir, with eco- 
logical notes. Biologia (Lahore) 25:37-50. 

Khan, M. S. 1980 Affinities and Zoogeography of her- 
petics of Pakistan. Biologia (Lahore), 26: 1 1 3- 1 7 1 . 

Khan, M. S. 1982a. Key for the identification of 
amphibian tadpoles from the plains of Pakistan. Paki- 
stan Journal of Zoology 14:133-145. 

Khan, M. S. 1982b. Collection, preservation and iden- 
tification of amphibian eggs from the plains of Paki- 
stan. Pakistan Journal of Zoology 14:241-243. 

Khan, M. S. 1991. Morphoanatomical specializations 
of the buccopharyngeal region of the anuran larvae 
and its bearing on the mode of larval feeding. Ph.D. 
thesis. Department Zoology, Punjab University, 
Lahore. 93 pp. 

Khan, M. S., and S. A. Malik. 1987a. Buccopharyn- 
geal morpology of tadpole larva of Rana hazarensis 
Dubois and Khan 1979, and its torrenticole adapta- 
tions. Biologia (Lahore), 33:45-60. 

Khan, M. S., and S. A. Malik. 1987b. Reproductive 
strategies in a subtropical anuran population in arid 
Punjab, Pakistan. Biologia (Lahore) 33:279-303. 
Khan. M. S., and S. A. Mufti. 1994a. Oral disc mor- 
phology of amphibian tadpole and its functional cor- 
relates. Pakistan Journal of Zoology 26:25-30. 

Khan, M. S., and S. A. Mufti. 1994b. Buccopharyn- 
geal specializations of tadpole of Bufo stomaticus and 
its ecological correlates. Pakistan Journal of Zoology 
26:285-292. 

Khan, M. S., and S. A. Mufti. 1995. Oropharyngeal 
morphology of ditritivorous tadpole of Rana cyanoph- 
lyctis Schneider, and its ecological correlates. Paki- 
stan Journal of Zoology 27:43-49. 

Khan, M. S., and R. Tasnim. 1987. A field guide to the 
identification of herps of Pakistan. Part I: Amphibia. 
Monograph No. 14. Biological Society of Pakistan 
(Lahore), 28 pp. 

Liu, C. C. 1950. Amphibians of western China. Field- 
iana: Zoology, Memoirs 2: 1-400. 



Vol. 9, p. 138 



Asiatic Herpetological Research 



2000 



Nishikawa, K., and R. Wassersug. 1988. Morphology 
of the caudal spinal cord in Rana (Ranidae) and Xeno- 
pus (Pipidae) tadpoles. Journal of Comparative Neu- 
rology 269:193-202. 

Nishikawa, K., and R. Wassersug. 1989. Evolution of 
spinal nerve number in anuran larvae. Brain Behav- 
iour Evolution 33: 1 5-24. 

Orton, G. L. 1953. The systematics of vertebrate lar- 
vae. Systematic Zoology 2:63-75. 

Rao, C. R. N. 1917. On the occurance of iridiocytes in 
the larva of Microhyla omata Boul. Record Indian 
Museum 13:282-292. 

Seale, D. B., and R. J. Wassersug. 1979. Suspension 
feeding dynamics of anuran larvae related to their 
functional morphology. Oecologia (Berlin) 39:259- 

272. 

Wassersug, R. J. 1980. The internal oral features of 
larvae from eight anuran families: Functional, system- 
atic, evolutionary, and ecological considerations. Uni- 
versity of Kansas Museum of Natural History, 
Miscellaneous Publication 68:1-146. 



Wassersug, R. J. 1984. The Pseudohemisus tadpole: a 
morphological link between microhylid (Orton type 
2) and ranoid (Orton type 4) larvae. Herpetologica 40: 
138-149. 

Wassersug, R. J. 1989. What, if anything is a micro- 
hylid (Orton type II) tadpole? Pp. 534-538. In H. 
Splechtna and H. Hilgers (eds.), Trends in vertebrate 
morphology. Progress in Zoology, 35. Gustav Fischer 
Verlag, Stuttgart and New York. 

Wassersug, R. J., and A. M. Murphy. 1987. Aerial res- 
piration facilitates growth in suspension feeding anu- 
ran larvae (Xenopus laevis). Experimental Biology 
46:141-147. 

Wassersug, R. J., and W. F. Pyburn. 1987. The biology 
of the Peret toad, Otopryne robusta (Microhylidae), 
with special consideration of its fossorial larva and 
systematic relation ships. Zoology Journal of Lin- 
naeus Society 9 1 : 1 37- 1 69. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 139-141 



Variation in Pelobates syriacus of Turkey 

Ismail H. UGurta§ 

Science and Art Faculty, Uludag University, Department of Biology, Bursa, Turkey 

Abstract.- Morphometric data and color patterns of Pelobates syriacus from different regions of Turkey were 
compared. Pelobates syriacus from Edirne appear to be distinct in terms of color pattern. Pelobates syriacus from 
Seydi§ehir appear to be distinct in terms of morphometric measurements. 

Key words.- Pelobates syriacus, Turkey, morphometry, color pattern 



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Figure 1 .The places where Pelobates syr/acushave been collected in Turkey. The key to the letters is found in 
Table 2. Dark circles = The localities where we collected specimens. Open circles = The localities where speci- 
mens were collected before. 1. Buyukdolluk, 2. Babaeski, 3. Altin§ehir, 4. Terkos, 5.Yassioren, 6. Adapazari, 7. 
Sogutlu, 8. Karasu, 9. Poyrazli Lake, 10. £erkesji, H.Terme, 12. Karacabey, 13. Bursa, 14. §akran, 15. Bostanli, 
16 Bornova, 17. Ivrindi, 18. Seydi§ehir,19. Karata§, 20. Iskenderun, 21. Van. 



Pelobates syriacus Boettger 1889 was first described 
from specimens collected from Hayfa-Israel. Later, 
Mertens (1923) examined two specimens, found in 
Belesuwar near the Azerbaijan and Iran border. 
Because of their long and narrow skull and the round 
and raised frontoparietals, he recognised them as 
another subspecies, Pelobates syriacus boettgeri. 

Other specimens from Macedonia were described 
by Karaman (1928) as the subspecies Pelobates syria- 
cus balcanicus. Miiller (1932) criticized this classifi- 
cation based on the shape of the skull. He emphasised 
that skull shape can be variable. Another subspecies 



living around Tiflis, Georgia was named Pelobates 
syriacus transcaucasicus by Delwig (1927). Accord- 
ing to Gilsen (1937), this subspecies is intermediate in 
morphology between P. s. boettgeri and P. s. syriacus, 
and Terentjev and Chernov (1965) are convinced that 
P. s. transcaucasicus is a synonym of the nominate 
race. Furthermore, Eiselt and Schmidtler (1973) pro- 
posed P. s. boettgeri as a synonym for P. s. syriacus. 

The taxonomic status of Turkish Pelobates syria- 
cus is poorly studied. Mertens (1953) described a 
young sample collected in Van (Turkey) and sug- 
gested that it was P. s. boettgeri. Zaloglu (1964) stud- 



Vol. 9, p. 140 



Asiatic Herpetological Research 



2001 



Table 1. Percentage of each color pattern in populations of Pelobates syriacus. 



Population 



B 



D 



Edirne 

Istanbul 

Adapazari 

Samsun 

Bursa 

Balikesir-izmir 

Seydi§ehir 

Adana-lskenderun 

Van 



- 


- 


100 


- 


56.66 


40.00 


3.34 


- 


11.54 


11.54 


11.54 


65.38 


68.42 


21.05 


- 


10.53 


73.92 


21.73 


- 


4.35 


15.00 


75.00 


5.00 


5.00 


7.70 


- 


- 


92.30 


66.00 


- 


- 


34.00 


14.38 


14.28 


. 


71.44 



ied Pelobates syriacus from the Turkish region of 
Izmir. He made an osteological comparison of chra- 
cters used to differntiate the subspecies of P. syriacus. 
He pointed out that these characters showed a wide 
range of variation, and refrained from classifying the 
Izmir P. syriacus into subspecies. 

In this study, samples (77 male, 57 female, 25 
juveniles, 35 larvae) collected from 9 different areas 
(Istanbul, Adapazari, Samsun, Bursa, Izmir-Balikesir, 
Seydi§ehir, Adana, Van) were investigated to obtain 
more information about variation in Pelobates syria- 
cus living in Turkey (Fig. 1 ). 

Results and Discussion 

There are four kinds of dorsal patterns on Pelobates 
syriacus (Fig. 2). The dorsal patterns are not sexually 
dimorphic. The percentage of dorsal patterns are fol- 
lows: A: 36.08 %, B: 25.95 %, C: 13.92 %, D: 24.05 
%. When all the samples are compared, it is seen that 
the dorsal pattern of Edirne population is different 
than the other populations (Table 1 ). 

All of the adults from Edirne have big green dots 
narrow raised surrounding on the back. The back- 
ground color of the back is light yellow-green. On this 
background there are connected raised surrounding 
with brown-green dots. Those dots also appear on all 
extremities. Furthermore, on the background color, 
and within the brown-green dots, are yellow and red 
dots. These dots are missing on the upper side of the 
front fingers. Dots on the body become smaller on the 
side. All the Edirne population have C pattern type 
(Fig. 2). 



In terms of morphological measurements, Pelo- 
bates syriacus populations living in Turkey are very 
similar. Here, the exception is the Seydi§ehir popula- 
tion which have shorter indices of tibia length (Table 
2). 

The differences in the color patterns of adult ani- 
mals from the Edirne population and the morphomet- 





Figure 2. Types of color patterns in Pelobates syria- 
cus. A: Spots are irregularly distributed and isolated. 
B: Two or more spots combine to form irregular islets. 
C: The edges of the spots are wavy and connected by 
thin bands. D: Spots form lengthwise bands. 



2001 Asiatic Herpetological Research Vol. 9. p. 141 



ric differences of the Seydi§ehir population should be 
explored using biochemical data in order to determine 
whether the Pelobates syriacus from these regions 
represent different taxa. 

Literature Cited 

Boettger, O. 1889. Ein neuer Pelobates aus Syrien. 
ZoologischerAnzeiger 12:144, 1889. 

Delwig, W. 1927. Eine neue Art der Gattung Pelo- 
bates Wagler aus dem zentralen Transcaucasus. Zool- 
ogischer Anzeiger 75, Heft 1/2:24. 

Eiselt, J. and J. F. Schmidtler 1973. Froschlurche aus 
dem Iran unter Beriick chtigung ausser-iranischer 
Populationgruppen. Annalen Naturhistorischen Muse- 
ums in Wien 77:181-243. 

Gislen, T. 1937. On the history of evolution and distri- 
bution of the European pelobatids. Zoogeographica 
3:119. 

Karaman, S. 1928. Contribution a 1' Herpetologie de 
Jugoslavja. Bulletin de la Societe Scientifique de Sko- 
plje 4. 2:129. 

Mertens, R. 1923. Beitrage zur Kenntnis der Gattung 
Pelobates Wagler Senckenbergiana 5(3/4): 1 1 8- 1 28. 

Mertens, R. 1953. Weiters zur Kenntnis der Herpeto- 
fauna der Asiatischen Tiirkei. Istanbul Univeritesi 
Fakultesi Mecmausi Seri B18: 373-375. 

Muller, L. 1932. Beitrage zur Herpetologie der 
Sudosteuropaischen Halbinsel. Teil I, Zoologischer 
Anzeiger 100, Heft 11/12:301-309. 

Teretnjev, P. and S. A. Chernov. 1965. Key to 
Amphibians and Reptiles. Israel Program for Scien- 
tific Translations, Jerusalem. 

Zaloglu, §. 1964. Ege bolgesinde bulunan Pelobates 
syriacus'un morfoloji, osteoloji ve biyolojisi ile ilgili 
ara§tirmalar. Ege Univeritesi. Fen Fakultesi ilmi Rap. 
Ser. No. 16:1-50. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 142-144 



Translation: A New Species of the Turtle Genus Ctav#(Testudoformes: 

Testudinidae) 

Ming-tao Song 

Shaanxi Institute of Zoology, Xi 'an Province, China 
(Current Address: Northern west Institute of Endangered Animals, Xi'an, China 710032) 

Original English Abstract.- Cuora pani, sp. nov. (Figs. 1, 2). Holotype, SIZ 80170, an adult male. Allotype, SIZ 
80171, an adult female. All of these type specimens were collected from Xujiaba (alt. 420 m) of Pingli County in 
Shaanxi Province, on June 17, 1981 by the author, and are preserved in the Shaanxi Institute of Zoology. This 
new species is similar to Cuora yurmanensis (Boulenger), but differs from the latter in having the median keel 
not conspicuous and no lateral keels; a brown narrow band extending from behind eye to the neck; the suture 
between gulars 1.5 times as long as that between humerals and much shorter than those between pectorals and 
between anals, but longer than that between femorals; the plastron yellow, with black sutures; and the limbs 
brown, without any markings. 

Song, M. T. 1984. A New Species Of The Turtle Genus Cuora (Testudoformes: Testudinidae). Acta Zootaxonom- 
ica Sinica 9(3):330-332. (In Chinese with English abstract) 



There are six species belonging to the genus Cuora 
Gray, 1855 (Pritchard, 1967; Sichuan Institute of 
Biology, 1977), that are distributed in Thailand, Cam- 
bodia, Malaysia, Indonesia, the Philippines, and 
southern provinces of China. 

In 1981, two Cuora specimens were found during a 
survey of Mt. Dabashan, Shaanxi Province. These 
specimens are different from the six known species 
and are recognized as a new form. Its description is 
given below. 

Cuora pani, sp. nov. (figs. 1-2) 

The holotype, SIZ 80170, is an adult male. The allo- 
type, SIZ 80171, is an adult female. Both of the type 
specimens were collected from Xujiaba (alt. 420 m) 
of Pingli County in Shaanxi Province, on June 17, 
1981 by the author, and are preserved in the Shaanxi 
Institute of Zoology. 

Diagnosis 

The new species is similar to Cuora yurmanensis 
(Boulenger), but differs from that species by having a 
median keel that is not conspicuous and no lateral 
keels; a brown narrow band extending from behind 
the eye to the neck; the seam between the gulars 1.5 
times as long as the seam between the humerals and 
much shorter than those between pectorals and 
between anals, but longer than that between femorals; 
the plastron yellow, with black markings along the 
seams; the limbs are brown and without any mark- 
ings. 



Description of Holotype 

The carapace is flattened, with an inconspicuous 
median keel. The cervical scute is small. The anterior 
margin of the first vertebral is flaring, the anterior 
margin wider than the posterior margin. The second 
vertebral is square-shaped, with both sides are slightly 
protruding. The third and the fourth vertebrals are as 
wide as long, with the fourth vertebral being wider 
than long. The first pleural is the longest, the second is 
wider and as long as the third, and the fourth the 
smallest. There are twelve marginals. The first mar- 
ginal is the widest, the third to seventh and the ninth 
to tenth are slightly flared. 

The plastron is rounded anteriorly and notched 
posteriorly. The plastron can completely close the 
shell and is united to the carapace by ligamentous tis- 
sue. The ligamentous tissue is also between the pecto- 
rals and abdominals. The length of gular seam is 1.5 
times the humeral seam, and much shorter than pecto- 
ral, abdominal, or anal seam. The pectoral seam is as 
long as abdominal seam, the anal seam a little shorter, 
and the abdominal seam is nearly twice the length of 
the gular seam; femoral seam much shorter, only a lit- 
tle longer than humeral seam. There are a pair of anals 
that are notched posteriorly. 

The head is moderate in size, smooth on top, and 
rather rough in the occipital region. The snout is 
pointed, projecting over the upper jaw. The diameter 
of the orbit is about the same of the length of snout. 
Upper jaw is slightly curved and a little longer than 
the lower jaw. 



2001 



Asiatic Herpetological Research 



Vol. 9. p. 143 





| = 1 cm 



I 



cm 



Figurel. The holotype (SIZ 80170), an adult male, in 
dorsal view. 



Figure 2. The allotype (SIZ 801 70), an adult female, in 
ventral view. 



The forelimb is covered by imbricate scales poste- 
riorly and with a transverse series of large scales on 
ventrally. The hindlimb is covered by scales medially 
and on the tarsus. There are five claws on the forelimb 
and four on the hindlimb. The webbing between the 
digits is well developed. The tail is short, conical in 
shape. The tail is covered by granular scales on its 
back, and covered with paired scales forming a longi- 
tudinal groove ventrally. 



Color in life 

The carapace is light brown. The plastron and ventral 
side of marginals are yellow, with broad black bars 
along the seams. The top of the head is olive, dark 
gray laterally, with two brown stripes behind the orbit 
and along tympanum to neck. The tympanum is light 
gray. The lower jaw and chin are grayish-yellow. The 
back of the neck is brown, but lighter ventrally. The 
shoulders are light yellow and the axilla are yellow. 
The limbs and tail are light brown above and gray 
below. The pelvic region and the area behind the fern- 



Vol. 9. p. 144 



Asiatic Herpetological Research 



2001 



ora are light yellow. The claws are brown with their 
tips yellow. 

Allotype 

The seams of the plastron are not as black as in the 
holotype, the back of tail is covered with a few large 
scales. 

Acknowledgments 

This translation was provided by Ermi Zhao with per- 
mission of Acta Zootaxonomica Sinica. The figures 
were provided by Ming-Tao Song. 

Table 1 . Measurements of types (in mm). 



Literature Cited 

Department of Herpetology. Sichuan Institute of Biol- 
ogy (Zhao. E.. Y.-m. Jiang, and Y. Shen). 1977. [Sys- 
tematic Key to Chinese Reptilia]. Science Press, 
Beijing. 1 10 pp. (in Chinese). 

Pope. C. H. 1935. The reptiles of China. Natural His- 
tory of Central Asia 10:28-35. 

Smith. M.A. The Fauna of British India. Including 
Ceylon and Burma. Reptilia and Amphibia. Vol. I.- 
Loricata, Testudines. Taylor and Francis. London. 185 
pp. 











































































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CD 

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a. 


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Holotype Male 32 18 16 18 120 86 40 

Allotype Female 33 18 18 27 115 81 37 



33.3 



32.2 



2000 



Asiatic Herpetological Research 



Vol. 9, pp. 145-148 



Translation: Testudo graeca anamurensis ssp. nov. from Asia Minor 

Heinz Weissinger 

Richard-Gebhardtgasse 4. 3423 St. Andra-Wordern, Austria 



Weissinger. H. 1987. Testudo graeca anamurensis ssp. nov. aus Kleinasien. OGH-Naehriehten (Wien) 10/1 
14-18 

The South-western coast of Turkey, from the Bey 
mountains near Antalya eastwards up to Mersin is the 
area of distribution of a Testudo graeca which is char- 
acterized hy an elongated, trapezium-shaped and 
strongly flattened carapace. This southcoast-popula- 
tion was so conspicuous among the more than 200 
specimens which I examined and measured in Turkey, 
that I consider them a geographic subspecies and 
describe them as follows: 



Testudo graeca anamurensis nov. ssp. 

Material 

Type: One female. Anamurum; NMW 30795:1 
Paratypes: One male, Anamurum; NMW 30795:2df 
One female. Side; NMW 30795:3 
Both alive in the author's collection 

Terra typica 

Beach of Anamurum. 7 km. west of Anamur. SW 
coast of Turkey. 

Diagnosis 

Differs from Testudo graeca ibera Pallas 1814 by 
its considerably Hatter and narrower (elongated trape- 
zium-shaped) carapace. The length of the carapace is 
two-and-a-half times its height and one-and-a-half 
times its width. In most cases the carapace pattern is a 
completely broken, obvious blotchy pattern. The plas- 
tron has a sharply demarcated pattern of brown to 
black blotches, which can cover the plastral area for 
20 - 100 7c. In 80 C A of cases, 100% at the terra typica, 
the animals have a white-yellow chin and throat. The 
head can be uniformly dark to irregularly spotted with 
yellow. A triangle on the snout - as in Testudo graeca 
terrestris Forskal 1775 - may be present only in early 
juvenile stages. In the subspecies most abundant in 
Asia Minor. Testudo graeca ibera Pallas 1814. the 
carapace is oval, domed regularly and has a smooth 
posterior margin. Its ground colour is bright olive with 
large yet individual blotches. The head and extremi- 
ties are most often uniformly dark. The plastron usu- 
ally shows indistinctly bordered blotches. 




Figures 1-3. (Top and bottom left) Testudo graeca 
anamurensis ssp. nov.; paratype. female NMW 
30795:3. 

Figure 4. (Bottom right) Testudo gracea. Ssp. anamu- 
rensis, male. Anamurum (left); ssp. ibera, female, 
Koyegegiz (right). 



Table 1. Measurements (straight-line distance) of the 
type specimen and a paratype 

NMW 30795:1 NMW 30795:3 



210mm 



Carapace 


260 mm 


length 




Carapace 
width 


160 mm 


Carapace 


105 mm 


height 





135 mm 



88 mm 



Vol. 9, p. 146 



Asiatic Herpetological Research 



2000 




303 tan 



9 Testudo graeca ibera 

C) Testudo graeca ibera " lerrestris " 



O Testudo graeca " lerrestris " 

J^ Testudo graeca anamurensis ssp. nov. 



Figure 5. Localities of Testudo graeca in Turkey with dates that tortoises were observed (Weissinger, 1 986). 1 , ca. 
70 km NW of Ankara, 5/18; 2, Dogansehir, open oak forest, 5/20; 3, 4 km from Diyabakir, 5/22; 4, 40 km from Mar- 
din; 5, Birecik, 5/24; 6, Nizip, in the direction of Ganziantep, dry wadi, about 45 g steep loam walls with tortoise bur- 
rows, 5/24; 7, ca. 10 km N of Islaniye, 5/24; 8, 20 km S of Iskenderund, 5/25; 9, ca. 20 km N. of Iskenderund, 5/26; 
10, ca. 25 km of Fg, cornfield, 5/26; 1 1 , 1 km from road crossing Ceyhan - Gaziantep, lava field, 5/26; 12, Ayas, 5/ 
27; 13, Korykos-Kizkale, in the direction of Silifke, 5/27; 14, Bozyazi, onion field, 5/28; 15, Anamurum, 7 km W of 
Anamur, 5/29; 16, ca. 20 km W of Anamur, 5/29; 17Pine forest, ca. 40 km W of Anamur, towards Gazipasa, 5/29; 
18, Side, sand dunes, ruin-meadows, 5/30; 19, 20 km from Antalya, 5/31; 20, 10 km from Antalya, towards Kugla, 
5/31; 21, Kemer, 5/31; 22, 10 km from Kemer, 5/31; 23, Mountain pass, ca. 800 m above sea level, towards Kas, 
5/31; 24, Vavi, ca. 800 m above sea level, towards Kas, 5/31; 25, Kinis, wheat field, 6/1; 26, Letoon, tilled field, 6/ 
1 ; 27, Esen, pine forest, 6/1 ; 28, Kargi, 6/1 ; 29, Koygegiz, riverine forest with wet meadows 6/2; 30, Ula, 6/2; 31 , 
Gokovalskele, 6/2; 32, Yatagan, 6/2; 33, Soke, towards Kusadasi, 6/2; 34, Selcuk, 6/3; 35, Pamucak, 6/4; 36, 2 km 
from Ayvacik, 6/6; 37, 15 km from Ayvacik, 6/6; 38, 20 km from Kesan, towards Greece (European Turkey). 



Etymology 

I name this South-west coast race after its terra typica. 
Anamurum, Testudo graeca anamurensis. 

Distribution 

The area of distribution of this race extends along 
the Southwest coast of Turkey from the Bey Daglari 
in the West to the plain of Mersin in the East. Most 
were found in the area between Side and Anamurum. 

Acknowledgments 

This translation was provided by Peter Paul van Dijk 
with some additions by Heinz Grillitsch. The transla- 
tion was done with the permission and full agreement 
of the Austrian Herpetological Society. Figures 1-4 
were provided by Jim Buskirk who, in turn, recieved 



Weissinger's original slides from his widow. The edi- 
tors of AHR would like to thank Heinz Grillitsch for 
his assistance in publishing this translation. 

Literature Cited 

Bodenheimer, F. S. 1935. Animal life in Palestine. An 
introduction to the problems of animal ecology and 
zoogeography. Ludwig Mayer, Jerusalem. 506 pp. 

Eiselt. T J., and Spitzenberger, F. 1967. Ergebnisse 
Zoologischer Sammelreisen in derTiirkei: Testudines. 
(Results of zoological collecting expeditions in Tur- 
key : Testudines) - Annalen des Naturhistorischen 
Museums in Wien. Serie B. fur Botanik und Zoologie 
70: 357-378. 



2000 



Asiatic Herpetologia.il Research 



Vol.9, p. 147 




Figure 6. The type specimen of Testudo graeca ana- 
/ra//-<?/75/5Weissinger 1987, NMW 30795:1. 



Figure 7. A male paratype of Testudo graeca anamu- 
/ms/sWeissinger 1987, NMW 30795:2. 



Obst. F.-J.. and Meusel. W. 1974. Die Landschild- 
kroten Europas (The tortoises of Europe) A. Ziemsen 
Verlag. Wittenberg Lutherstadt 

Wermuth. H. 1958. Status und Nomenklatur der Mau- 
rischen Landschildkrote, Testudo graeca, in SW- 
Asien und NO-Afrika. (Status & Nomenclature of T. 
graeca in SW Asia & NE Africa). Senckenbergiana 
Biologica. 39 : 149-153. 



Wermuth. H.. and Mertens. R. 1961. Schildkroten - 
Krokodile - Briickenechsen. (Turtles. Crocodiles & 
Tuatara). Fischer Verlag. Jena 422 pp. 

Werner, F 1902. Die Reptilien- und Amphibienfauna 
von Kleinasien. (The reptile and amphibian fauna of 
Asia Minor). Sit/^ungsberichte Akademie Wissen- 
schaften. Mathematisch-Naturwissenschaftliche 

Klasse 111: 1057-1121. 



Vol. 9, p. 148 



Asiatic Herpetological Research 



2000 



Appendix 

This translation was prepared by Peter Paul van Dijk 
with some additions by Heinz Grillitsch of the 
Naturhistorische Museum Wien (NMW). This trans- 
lation is published with the full permission of the 
Austrian Herpetological Society. In addition to the 
original figures. Heinz Grillistach and Alice Schuma- 
cher (NMW) have provided additional figures of the 
original type series (Figs. 6-8). Uwe Fritz (pers. 
comm.). states that there are three additional 
paratypes in the collections of the Staatliches 
Museum fur Tierkunde Dresden (MTKD 29200- 
29202). Finally, the female paratype that Weisssinger 
lists as NMW 30795 is currently catalogued as NMW 
31031 (Fritz and Grillitsch, pers. comm). 




Figure 8. A female paratype of Testudo graeca anamu- 
/•ews/sWeissinger 1987, NMW 31031 (formerly 
30795:3). 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 149-150 



Book Review: A Guide to the Fauna of Iran 



A Guide to the Fauna of Iran. Eskandar Firouz. Iran University Press; Tehran. 2000. vi + 491 pp 



The first attempt to provide a comprehensive verte- 
brate zoology of Iran was that of William T. Blanford 
in 1876. Since that time, there have been treatises on 
various groups of vertebrates (e. g., mammals: Lay. 
1967; Harrington, 1977; Ziaie, 1996; birds: Scott, et 
al., 1975; reptiles: Latifi, 1984 (snakes); Anderson, 
1979 (turtles, crocodiles, and amphisbaenians), 1999 
(lizards); amphibians: Baloutch and Kami, 1995; 
fishes: Coad, 1987, 1995. A Guide to the Fauna of 
Iran by Firouz is the first attempt since Blanford to 
bring all of the vertebrates of Iran together into a natu- 
ral history guide. 

Although the text is in Farsi (Persian), it is acces- 
sible to Western readers as a comprehensive list of the 
vertebrate species of Iran, since there is a Latin and 
English index, color illustrations with Latin species 
names, and lists of taxa with both Latin species names 
and English colloquial names. Most genera are repre- 
sented by color illustrations, fishes by paintings, 
amphibians and reptiles by photographs, birds by 
paintings, and mammals by both. A list of the princi- 
pal works consulted is provided (pp. 423-432). This 
list constitutes a good beginning bibliography for any- 
one interested in the vertebrate zoology and natural 
history of Iran. 

For this book, the author has consulted the zoolo- 
gists currently working on each of the vertebrate 
groups to assure that the species lists are up to date. 
The work covers 164 vertebrate families and 1054 
species. Emphasis is given to conservation topics, 
including threatened and endangered species, prob- 
lems of exotic introductions, and the ecological con- 
sequences of environmental change. The natural 
history sections include consideration of the zoogeog- 
raphy of Iran. 

A renewed interest in zoology has occurred 
recently in Iran, the many universities including 
departments of biology and zoology are attempting to 
build knowledgeable faculties in these disciplines. 
This emphasis is largely due to the efforts of the 
author, Eskandar Firouz. Prior to the Islamic Revolu- 
tion, Firouz was the principal architect of the conser- 
vation, natural history legislation and implementation 
in Iran through the Department of Environment, the 
establishment of Protected Regions and National 
Parks, and the regulation of hunting and fishing. 
These efforts were well on the way to setting a world 
standard for conservation and the study of natural his- 
tory at the time of the revolution. Interests in natural 



history have survived the period of resistance to per- 
ceived "Westernization" in Iran, and young scholars 
and conservationists must now face the daunting chal- 
lenge of rebuilding the edifice of conservation initi- 
ated by Firouz and his colleagues in the 1960s and 
early 70s. The present book is yet another major con- 
tribution to that effort, and there should be a well-used 
copy in every local Department of Environment 
office, university and department library and in the 
personal libraries of individual zoologists and ecolo- 
gists in Iran. It will also be a useful reference in spe- 
cialized libraries, natural history museums, and 
conservation institutions in the West and in the coun- 
tries of the former Soviet Union. 

Steven C. Anderson 

Department of Biological Sciences, University of the 

Pacific, Stockton, CA 95211 USA and Department of 

Herpetology, California Academy of Sciences, Golden 

Gate Park, San Francisco, CA, 94118 USA. email: 

asaccus@aol. com 

Literature Cited 

Anderson, S. C. 1979. Synopsis of the turtles, lizards, 
and amphisbaenians of Iran. Proceedings of the Cali- 
fornia Academy of Sciences, ser. 4, 41(22):501-528. 

Anderson. S. C. 1999. The Lizards of Iran. Society for 
the Study of Amphibians and Reptiles, Ithaca, NY, vii 
+ 442 pp. 

Baloutch, M. and H. G. Kami. 1995. Amphibians of 
Iran. Tehran University Publications, Tehran, 177 pp. 
(In Farsi [Persian]). 

Blanford, W. T. 1876. Eastern Persia. An Account of 
the Journeys of the Persian Boundary Commission, 
1870-1872, vol. 2. The Zoology and Geology. Mac- 
millan and Co., London, viii + 516 pp. 

Coad, B. W. 1987. Zoogeography of the freshwater 
fishes of Iran. In : Krupp, et al. (Eds.). Proceedings of 
the Symposium of the Fauna and Zoogeography of 
the Middle East. Beihefte zum Tubinger Atlas des 
Vorderen Orients. Reihe A (Naturwissenschaften) 
28:213-228. 

Coad, B. W. 1995. Freshwater fishes of Iran. A check- 
list. Institute of Landscape Ecology of the Academy 
of Science of the Czech Republic, 29( 1 ): 1 -64. 



Vol. 9, p. 150 



Asiatic Herpetological Research 



2001 



Harrington, F. A. Jr. 1977. A Guide to the Mammals 
of Iran. Department of the Environment, Tehran, 89 
pp. 

Latifi, M. 1984. The Snakes of Iran. Department of 
the Environment, Tehran, 231 pp. (in Farsi [Persian]; 
English Translation, 1991, Society for the Study of 
Amphibians and Reptiles, Oxford, Ohio). 

Lay, D. M. 1967. A study of the mammals of Iran. 
Fieldiana Zoology 54:282 pp. 



Scott, D. A., H. Moravej Hamadani, and A. Adhami 
Mirhosseyni. 1975. Parandegan-e Iran (The Birds of 
Iran). Department of the Environment, Tehran (In 
Farsi [Persian]; Latin, English, and French names). 

Ziaie, H. 1996. Field Guide to the Mammals of Iran. 
Department of the Environment, Tehran (In Farsi 
[Persian]; Latin and English names). 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 151-152 



Book Review: Four Recent Handbooks for Turkey 



Tiirkiye Amfibiliri. The Amphibians of Turkey by 

Necla Ozeti and I. Yilmaz, 1994. Izmir, 219 pp., 40 
col. photos in pis., 92 text-figs. (In Turkish with 
English summary). 

Tiikiye Omurglilari, Amfibiler edited by Ali Demir- 
soy, 1996. Meteksan; Ankara, vi + 69 pp., numerous 
maps and figs. (In Turkish). 

Tiikiye Omurglilari, Siiriingenler edited by Ali 
Demirsoy, 1996. Meteksan; Ankara, viii + 205 pp., 
numerous maps and figs. (In Turkish). 

Turkish Herpetofauna (Amphibians and Reptiles) 

by Ibrahim Baran and Mehmet Atatiir. 1998. Republic 
of Turkey Ministry of Environment: Ankara, x + 214 
pp., 122 col. pis., 7 text-figs. (In English). 

Until the publication of these books, the standard 
herpetological summary publications for Turkey had 
been Ba§oglu and Ozeti (1973) and Ba§oglu and 
Baran (1977, 1980). The four handbooks reviewed 
here accompanied us on a recent herpetological expe- 
dition to western and southern Turkey. The combina- 
tion of these references enabled us to quickly identify 
most species in the field. 

Turkish Herpetofauna by Baran and Atatiir has a 
color photograph of each species, illustrated keys, one 
paragraph each on identification, habitat, biology, and 
distribution. The distribution sections include brief 
statements of the overall distribution and the range 
within Turkey. Where subspecies are recognized, a 
short distribution statement is given for each. There is 
no descriptive differentiation of subspecies. There are 
no distribution maps. The color photographs are of 
good quality and usually enabled quick identification 
of live specimens. Photos and text are on glossy paper 
and this hardback book is sturdily bound. The bibliog- 
raphy includes 53 references, and while not exhaus- 
tive, serves as a good introduction to the 
herpetological literature of Turkey. The nomenclature 
is reasonably current, with the exception, of course, of 
changes (e.g. the break-up of Eumeces and Coluber) 
since its publication. It is the only one of these hand- 
books to include Rana dalmatina Schneider, Sinch, 
and Nevo, 1992. Of the four, this book got most use 
on the trip, particularly as it is in English. 

The two paper-bound books edited by Demirsoy, 
Tiikiye Omurglilari, Amfibiler and Tiikiye Omurg- 
lilari, Siiriingenler are useful to Western herpetolo- 
gists, in particular, because they have a distribution 
map for each species, usually accompanied by a black 
and white illustration, often a photograph, but some- 
times one or more line drawings. Although the books 



are printed on high quality stock, many of the photo- 
graphs have not reproduced well. 

The tabular format of species presentation is 
somewhat unusual and innovative. These tables are 
the same for each species and include Latin and Turk- 
ish names, the author, date; one or more illustrations; 
distribution map for Turkey, showing both range and 
spot localities; remarks on habitats, relationships, tax- 
onomic status, etc.; conservation status (rare, endan- 
gered, vulnerable, widespread, etc.); overall 
distribution and range within Turkey; type locality; 
greatest size; morphological characteristics; color pat- 
tern; reproduction; time from hatching to maturity; 
conservation status of habitats; defense mechanisms; 
population densities; potential hazards; zoogeo- 
graphic origins; and conservation proposals for the 
future. A table listing the fauna summarizes the global 
and Turkish distributions, zoogeographic source, pre- 
sumed geological time of arrival in Turkey, biogeo- 
graphic and conservation status, and special remarks. 

The books also contain descriptions and com- 
ments on higher taxa for the general reader. The bibli- 
ographies are extensive, much more so than that of the 
Baran and Atatiir handbook in English. Contributors 
to these volumes are prominent Turkish herpetolo- 
gists: Varol Tok, Ibrahim Baran, Mehmet Atatiir, Abi- 
din Budak, and Mehmet Oz. These volumes are 
obviously designed for the serious Turkish zoology 
student or wildlife professional and should serve this 
purpose well. They appear to be part of a larger series 
covering the entire Turkish fauna. Subspecies are cov- 
ered individually, although there are some strange 
inconsistencies: sometimes there is a separate table 
and map for the species as a whole, in addition to the 
subspecies accounts, and sometimes not. The usage of 
generic names is sometimes confusing, as for exam- 
ple: Cyrtodactylus (-Cyrtopodion) (Mediodactylus) 
heterocercus and Agama (-Laudakia) stellio so that it 
not clear which generic name is recognized by the 
authors. As I don't read Turkish, I can't comment on 
errors in the text. Some of the range maps may be too 
general, especially in those cases where spot distribu- 
tions are not shown. For example, Laudakia stellio is 
shown to extend to broad contact with the Iranian bor- 
der, whereas it has never been recorded for Iran. We 
used the maps of these volumes in conjunction with 
the photographs and English text of the Baran and 
Atatiir book and this greatly facilitated our work. 

Tiirkiye Amfibiliri. The Amphibians of Turkey by 

Ozeti and Yilmaz is aimed primarily at Turkish zool- 
ogy students and serves as an introduction to amphib- 



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Asiatic Herpetological Research 



2001 



iology. It is an update, or second edition of Ba§oglu 
and Ozeti (1973). The first chapter is an introduction 
to nomenclature, and in particular, to the taxonomy of 
the Turkish amphibian fauna. One chapter deals with 
the biology of amphibians in general, including mor- 
phology, reproduction, development, distribution, and 
folklore. Two subsequent chapters cover these topics 
in more detail, one for salamanders and one for frogs. 
These chapters are arranged by family and include 
families not found in Turkey. They include keys to the 
Turkish genera and species and spot locality maps. 
These maps are useful, if somewhat cluttered (several 
taxa per map); localities are numbered and named in 
the figure legends. Keys to the generic level for egg 
masses and larvae have been provided. The descrip- 
tions and discussions of morphology are accompanied 
by useful illustrations taken from several sources (not 
always attributed). There is a short chapter on meth- 
ods and techniques, a glossary, and a bibliography of 
181 references, including the most important techni- 
cal papers for the fauna, an excellent introduction to 
the literature of the Turkish amphibian fauna. At the 
end of the book are 1 3 plates of black and white pho- 
tographs illustrating the covered taxa and some habi- 
tats. The photographs (or at least the printing of them) 
are of rather poor quality, although they are adequate 
to illustrate pattern differences between the species. 
An English summary includes a comprehensive key to 
the amphibians of Turkey. This key, along with the 
English figure legends, and specific collecting locali- 
ties, makes this book particularly useful to non-Turk- 
ish speaking herpetologists. 



Steven C. Anderson 

Department of Biological Sciences, University of the 

Pacific, Stockton, CA 9521 1 USA and Department of 

Herpetology, California Academy of Sciences, Golden 

Gate Park, San Francisco, CA, 94118 USA. email: 

asaccus @ aol. com 

Literature Cited 

Ba§oglu, M. and N. Ozeti 1973. Tiirkiye Amfibileri. 
The Amphibians of Turkey. Ege Universitesi Fen 
Fakultesi Kitaplar Serisi No. 50. Ege Universitesi 
Matbaasi; Bornova/Izmir, 155 pp., 9 pis. 

Ba§oglu, M. and I. Baran. 1977. Tiirkiye 
Suriingenleri. Kisim I. Kaplumbaga ve Kertenkeleler. 
The Reptiles of Turkey. Part I. The Turtles and Liz- 
ards. Ege Universitesi Fen Fakultesi Kitaplar Serisi 
No. 76. Ilker Matbaasi; Bornova/Izmir, 255 pp., 16 
pis. 

Ba§oglu, M. and I. Baran. 1980. Tiirkiye 
Suriingenleri. Kisim II. Yilanlar.The Reptiles of Tur- 
key. Part II. Snakes. Ege Universitesi Fen Fakultesi 
Kitaplar Serisi No. 81. Ilker Matbaasi; Bornova/Izmir, 
218 pp. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 153-153 



Book Review: Wild About Reptiles. 
Field Guide to the Reptiles and Amphibians of the UAE 

Wild About Reptiles. Field Guide to the Reptiles and Amphibians of the UAE. by Marycke Jongbloed. 2000. 
Barkers Trident Communications; London. I 16 pp.. numerous unnumbered color photographs. Order from Natu- 
ral History Book Service: www.nhbs.com 



This attractive little spiral-bound book is the first field 
guide to the amphibians and reptiles of the United 
Arab Emirates. Most of the 68 species covered are 
illustrated with good color photographs. There are no 
keys, but the color photographs will serve to identify 
most live specimens. Descriptions are sketchy and the 
emphasis is on natural history. The book should serve 
well as an introduction to desert reptiles, sea snakes 
and sea turtles, and the few amphibians of the region. 
English speakers who work in the Emirates, particu- 
larly members of the natural history societies, will 
find this a handy reference. Visiting herpetologists. 
while they will still want the keys and descriptions of 
Arnold (1986). Balletto, et al. (1986). Gasperetti 
(1988). Leviton and Anderson (1967), and Leviton, et 
al. ( 1992). will want this for quick identification. The 
technical papers that emphasize the herpetology of the 
Emirates are few in number (see Arnold. 1984: Levi- 
ton and Anderson, ) and only Arnold ( 1984) concerns 
the ecology and natural history of this region of the 
Arabian Peninsula. 

Jongbloed takes a highly personal and anecdotal 
approach in this book, demonstrating her enthusiasm 
for these creatures. She includes field notes taken by 
her friend. John Norman Bishop "Bish" Brown. 
Because so little professional herpetological work has 
been done in the Emirates, and those studies that have 
been carried out have dealt mainly with systematics. 
using preserved specimens, this book is almost the 
only source of natural history information for these 
groups in the UAE. The natural history observations 
are informal, but still, informative. The bibliography 
is sketchy and disappointing and will not lead the 
beginning naturalist very deeply into the literature of 
the UAE. 

I recommend this field guide to any naturalist, pro- 
fessional or amateur, planning to travel to the UAE. 



Anyone seriously interested in the herpetofauna of 
Southwest Asia in general or the Arabian Peninsula in 
particular will want this in his library. 

Steven C. Anderson 

Department of Biological Sciences, University of the 

Pacific, Stockton, CA 9521] USA and Department of 

Herpetology, California Academy of Sciences, Golden 

Gate Park, San Francisco, CA, 94118 USA. email: 

asaccus @ aol. com 

Literature Cited 

Arnold. E. N. 1984. Ecology of lowland lizards in the 
eastern United Arab Emirates. Journal of Zoology, 
London 204:329-354. 

Arnold, E. N. 1986. A key and annotated check list to 
the lizards and amphisbaenians of Arabia. Fauna of 
Saudi Arabia 8:384-435. 

Balletto. E.. M. A. Cherchi, and J. Gasperetti. 1986. 
Amphibians of the Arabian Peninsula. Fauna of Saudi 
Arabia 7:318-392. 

Gasperetti. J. 1988. Snakes of Arabia. Fauna of Saudi 
Arabia 9:169-450. 

Leviton, A. E. and S. C. Anderson. 1967. Survey of 
the Sheikhdom of Abu Dhabi. Arabian Peninsula. Part 
II: Systematic account of the collection of reptiles 
made in the Sheikhdom of Abu Dhabi by John Gas- 
peretti. Proceedings of the California Academy of Sci- 
ences (4) 35(91:157-192. 

Leviton, A. E.. S. C. Anderson. K. Adler, and S. A. 
Minton. 1992. Handbook to Middle East Amphibians 
and Reptiles. Society for the Study of Amphibians 
and Reptiles. Oxford. Ohio, vii + 252 pp. 



2001 



Asiatic Herpetological Research 



Vol. 9, pp. 154-155 



Obituary: Sherman Anthony Minton Jr. 

Muhammad S. Khan 

7740 NW, 47th Street Lauderhill, FL.33351, USA 



Dr. Sherman Anthony Minton Jr., died of cancer in 
Indianapolis. Indiana, USA. on 15 June 1999, at the 
age of 80. He was born on the 24 February 1919, in 
New Albany. Indiana, where he also attended school. 
He had a fondness for amphibians and reptiles even in 
his childhood. He was the eldest child of Senator 
Sherman Minton Sr.. on whose suggestion young 
Sherman chose medicine instead of law, since medi- 
cine was closer to zoology. He obtained a BS in Zool- 
ogy in 1939 and an MD in 1942 at Indiana University. 
He served during World War II in the U.S. Navy from 
1943-1946. After the war, Sherman spent 1947-1948 
at the University of Michigan Zoology Department, 
taking Herpetology and Microbiology. He later joined 
the faculty of Indiana University School of Medicine, 
in the Department of Microbiology and Immunology, 
where he remained until he retired as Emeritus Pro- 
fessor in 1984. Dr. Minton's career presents a beauti- 
ful amalgamation of medicine, microbiology, and 
herpetology. 

Sherman married Madge Alice Shortridge Ruther- 
ford on 10 October 1943. while both were in military 
service. He was in the Navy and she was flying with 
the Women Air Force Service Pilots (WASP). Interest- 
ingly. Madge has also been interested in snakes from 
her childhood, so their relationship was also profes- 
sional. They co-authored eight publications including 
two books. Venomous Reptiles (Minton and Minton 
1969. 1980) and Giant Reptiles (Minton and Minton. 
1973). 

Minton took a break from Indiana ( 1958-1962) to 
teach at the Basic Medical Sciences Institute. Karachi. 
Pakistan (now Postgraduate Medical Center), as part 
of the U.S. AID program. He taught at the institute, 
reorganized the medical teaching courses, and headed 
the institute for a short time. 

He pioneered herpetological studies in Pakistan. 
Both he and his wife Madge Minton traveled 44,000 
miles in different parts of Pakistan, collecting 
amphibians and reptiles and information about them 
from the local people. One of the results of this is the 
well-illustrated paper on the amphibians and reptiles 
of Sind and Las Bela (1962). In 1965. he visited Iran 
and Pakistan under the sponsorship of American 
Museum of Natural History and collected material for 
his main book on the herpetology of Pakistan ( 1966). 




Sherman, with Madge's constant support and help, 
produced over 170 articles, books and monographs. 
He was a pioneer in the study of venomous reptiles 
and toxicology. He was on the editorial board of the 
journals Toxicon and Clinical Toxicology. He was the 
chairman and a member of the major professional 
societies in his field and received numerous awards 
and honors for his work. He also made a significant 
contribution to the herpetology of Indiana 

From 1972-1980 he joined several expeditions to 
different oceans to study biology of sea snakes and 
other venomous sea animals. He was visiting profes- 
sor in the Department of Zoology, University of New 
England in Australia during 1980. 

In remembrance of Dr. Minton's services to the 
herpetology of Pakistan, the following fossil Ameri- 



2001 Asiatic Herpetological Research Vol. 9, p. 155 

can frog and Pakistani lizard and snakes have been 
named after him: 

Proacris mintoni Holman, 1961 

Coluber karelini mintonorum Mertens, 1969 

Gymnodactylus mintoni Golubev and Szczerbak, 
1981 

Typhlops madgemintonai shermanai Khan, 1999 

Dr. Sherman A. Minton, Jr., is survived by his 
widow Madge Rutherford Minton, and three daugh- 
ters. 

Literature Cited 

Golubev. M., and Szczerbak, N. N. 1981. A new spe- 
cies of Gymnodactylus Spix, 1923 (Reptilia, Sauria, 
Gekkonidae) from Pakistan. Vestnik Zoolii 1981:40- 
45, (in Russian). 

Holman, A. 1961. A fossil frog from the Lower 
Miocene of Florida. Copeia 1961(3):354-355. 

Khan, M. S. 1999. Two new species and a subspecies 
of blind snakes of genus Typhlops from Azad Kashmir 
and Punjab, Pakistan. Russian Journal of Herpetology 
6(3): 231-240. 

Mertens. R. 1969. Die Amphibien und Reptilien West- 
Pakistans. Stuttgarter Beitrage zur Naturkunde 197: 1- 
96. 

Minton, Jr., S. A. 1962. An annotated key to the 
amphibians and reptiles of Sind and Las Bela, West 
Pakistan. American Museum Novitates 208 1 : 1-21. 

Minton, Jr., S. A. 1966. A contribution to the herpetol- 
ogy of West Pakistan. Bulletin of the American 
Museum of Natural History 134(2): 31-184. 

Minton Jr., S. A., and Minton, M. R. 1969. Venomous 
Reptiles. Charles Scribner's Sons, New York. 274 pp. 

Minton Jr., S. A., and Minton, M. R. 1973. Giant Rep- 
tiles. Charles Scribner's Sons, New York. 345 pp. 

Minton Jr., S. A., and Minton, M. R. 1980. Venomous 
Reptiles. Charles Scribner's Sons, New York. 308 pp. 

Stewart, M. M. 2000. Madge and Sherman Minton. 
Copeia, 2000(1): 304-309. 



2001 Asiatic Herpetological Research Vol. 9, pp. 156-160 



Guidelines for Manuscript Preparation and Submission 

NOTE: AHR now requires all tables to be submitted camera-ready. Please send 
hard copy, or electronic images of tables in the following format: 

Table 1 .AHR Standard table format. Note double line under heading row. Please use a sanserif font 
(Helvetica 1 0pt or Ariel 10pt). 



Column 1 Column 2 Column 3 Column 4 



12.4 43321.5 9.1 0.01 

12.1 91020.4 0.6 0.02 



Summary 

Manuscripts must: 

1) be written in English. 

2) be of letter quality (laser printed or typewritten on bond paper). 

3) include camera ready tables and figures (if any). 

4) include complete and accurate literature citations. 

5) include complete and accurate localities with latitude and longitude. 

6) include a camera ready map illustrating regions discussed (when applicable). 

Tips for electronic submission 

• Do not type authors names in all capitals in literature cited. 

• Do not use two spaces following a period, or for any other purpose. 

• Do not attempt to recreate the format of the journal in your manuscript. Please use only sim- 
ple formatting limited to italics, boldface, and underline. 

Manuscripts failing to meet these criteria will be returned without review for correction. 

Purpose and Content 

Asiatic Herpetological Research publishes articles concerning but not limited to Asian herpetology. The editors 
encourage publications from all countries in an attempt to create an open forum for the discussion of Asian her- 
petological research. 
Articles should be in standard scientific format and style. The following sections should be included: 

Title 

The title should reflect the general content of the article in as few words as possible. The editors encourage titles 
that summarize the main findings of the article. 

Names and Addresses 

The names and addresses of all authors must be complete enough to allow postal correspondence. Please include 
email and World Wide Web addresses if applicable. 

Abstract 

The abstract should briefly summarize the nature of the research, its results, and the main conclusions. Abstracts 
should be less than 300 words. 



2001 Asiatic Herpetological Research Vol.9, p. 157 



Key Words 

Key words provide an index for the filing of articles. Key words provide the following information (when appli- 
cable): 1 ) Taxonomy (e.g. Reptilia, Squamata, Gekkonidae, Gekko gecko). 2) Geography (e.g. China. Thailand). 
3) Subject (e.g. taxonomic validity, ecology, biogeography). The order of taxonomy, geography, and subject 
should be observed. 

Text 

Manuscripts must be in English and spelling must be correct and consistent. Use Webster's New International 
Dictionary for reference. For clarity, use active voice whenever possible. For example, the following sentences in 
active voice are preferable to those in passive voice. 

Active voice: "Lizards were extremely common on the site." and "I examined three female snakes." 

Passive voice: "Lizards were observed to be extremely common on the site." and "Three female snakes were 
examined." 

Abbreviation 

Do not abbreviate unless the full phrase has already appeared. Scientific names may be abbreviated only if they 
have appeared fully in the same paragraph. Never begin a sentence with an abbreviation of a scientific name. 

Statistics 

Statistics must be accompanied by sample sizes, significance levels, and the names of any tests. Investigators 
should pay careful attention to independence and applicability of tests, and randomness of samples. One of the 
most frequent examples of nonindependence is the use of multiple, paired t-tests instead of analysis of variance 
(anova). In general, multiple tests on the same data set are not valid. Descriptive statistics are in many cases more 
appropriate than inferential statistics. 

Standard Format 

Manuscripts following standard format should include introduction, methods, results, and discussion sections. 
While other formats are acceptable, the editors encourage the use of standard format. Please do not type in all 
capital letters. 

Introduction 

The introduction typically states the significance of the topic and reviews prior research. 

Material and Methods 

This section should clearly state where, when, and how research was carried out. Include sample sizes. Protocols 
designed by other investigators must be properly cited. Research materials and their manufacturers should be 
listed. The reader must be able to replicate the methods of the author(s). 

Results 

This section states the results and their significance to the investigation. Figures and tables may be used to clarify, 
but not to replace, results statements in the text. Statistics should be used when applicable. Large amounts of data 
should be avoided, or included as an appendix at the end of the article. 

Discussion 

The discussion is a synthesis of the introduction and the results. No new information should be discussed unless 
it was presented in the results section. New findings should be discussed in relation to prior research. The 
author(s) should feel free to present several possible interpretations of the results. The editors particularly 
encourage suggestions of future research in Asian herpetology. 



Vol. 9, p. 158 Asiatic Herpetological Research 2001 

Manuscript Preparation 

Overview 

Please do not attempt to replicate the formatting style of AHR in your manuscript. All formatting except italics 
will be removed in the production process. Bold and underlined text should be used only to identify section head- 
ing levels (see below). Extraneous formatting is counterproductive and increases the production costs of the jour- 
nal. There are a few simple guidelines that authors must follow. 

Section Headings 

Articles will be published using three section heading styles. All heading levels must be on their own line, and 
left justified. For the purposes of manuscript submission. Level 1 heading is bold, and generally reserved for 
Introduction, Material and Methods, Results, and Discussion; Level 2 is italic, and Level 3 is underlined . 

Figures 

Figures must be referenced in order in the text. Each figure illustration (line art or photograph) submitted must 
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one figure to a sheet. AHR cannot be responsible for redrawing, touching up, or otherwise modifying figure illus- 
trations for authors. In addition, figure illustrations submitted must: 

1 ) be of publication quality with typeset text. 

2) be mounted on a separate 21.5 x 28 cm (8.5 x 1 1 inch) sheet with figure number on back. 

3) be on a separate sheet from figure legend. 

4) not have poor type or handwriting on the face of the figure. 

5) The TIFF file format is preferable for electronic versions of figures, but Photoshop, JPEG, or PICT formats are 
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Figure Legends. Figure legends should be typed on a separate sheet. Legends should explain the figure without 
reference to the text. A figure and legend should make sense if separated from the rest of the article. For example: 

Figure 2. Lateral view of live Psammodynastes pulverulentus holding a prey lizard (Anolis car- 
olinensis). Note buccal tissue surrounding the enlarged anterior maxillary and dentary teeth of 
the snake. 

Color Figures. AHR may publish color figures at the discretion of the editors. AHR is now published both on 
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Tables 

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Typeface 

Twelve point type is preferred. Supply a detailed list of special characters (greek letters, male or female symbols, 
etc.) that are not part of a standard font. 



2001 Asiatic Herpetological Research Vol. 9, p. 159 



Literature Cited 

Accurate and standard references are a crucial part of any article. This is especially important when dealing with 
publications from many different countries. The reader must be able to precisely identify any literature cited. 
References in the text must be checked for consistency with references in the literature cited section. All refer- 
ences cited in the text must be in the literature cited section. The literature cited section may not contain any ref- 
erences not mentioned in the text. Articles containing inaccurate or inconsistent literature citations will be 
returned for correction. 

References in Text. 1 ) References to articles by one or two authors must include both surnames in the order they 
appear in the original publication. References to articles by more than two authors must include the first author's 
surname, followed by "et al." 2) The year of article follows the authors, separated only by a space. 3) References 
with the same author and year are distinguished by the lower case characters "a, b, c, . . ." 4) References cited in 
text are listed in alphabetical order by first author. 

For example, "My results also incorporate literature records (Marx et al., 1982; Marx and Rabb, 1972; 
Mertens, 1930; Pope, 1929; Wall, 1909, 1910a, 1910b, 1910c)." 

References in Literature Cited. 1 ) References must include all authors, in the order that they appear in the orig- 
inal publication; "et al." is never used in a literature cited section. 2) The first author is listed surname first, ini- 
tial(s) last. All other authors are listed initial(s) first, surname last. 3) References with the same author and year 
are distinguished by the lower case characters, "a, b, c, . . ." 4) References cited are listed in alphabetical order by 
first author. 5) Names of journals are not abbreviated. See below for examples: 

Journal article 

Dial, B. E. 1987. Energetics and performance during nest emergence and the hatchling frenzy in loggerhead sea 
turtles (Caretta caretta). Herpetologica 43(3);307-315. 

Journal article from a journal that uses year instead of volume 

Gatten, R. E. Jr. 1974. Effect of nutritional state on the preferred body temperatures of turtles. Copeia 
1974(4):912-917. 

Journal article, title translated, article not in English 

Ananjeva, N. B. 1986. [On the validity of Megalochilus mystaceus (Pallas, 1776)]. Proceedings of the Zoological 
Institute, Leningrad 1 57:4- 1 3. (In Russian). 

Note that for Acta Herpetologica Sinica, the year must precede the volume number. This is to distinguish 
between the old and new series, and between 1982-1987, Vols. 1-6 (new series) and 1988 with no volume number, 
numbers 1 and 2 (new series). 

Cai, M., J. Zhang, and D. Lin. 1985. [Preliminary observation on the embryonic development of Hyiwbius chin- 
ensis Guenther]. Acta Herpetologica Sinica 1985, 4(2): 177- 180. (In Chinese). 

Book 

Pratt, A. E. 1892. To the snows of Tibet through China. Longmans, Green, and Co., London. 268 pp. 

Article in book 

Huey, R. B. 1982. Temperature, physiology, and the ecology of reptiles. Pp. 25-91. In C. Gans and F. H. Pough 
(eds.). Biology of the Reptilia, Vol. 12, Physiological Ecology. Academic Press, New York. 

Government publication 

United States Environmental Data Service. 1968. Climatic Atlas of the United States. Environmental Data Ser- 
vice, Washington, D. C. 

Abstract of oral presentation 

Arnold. S. J. 1982. Are scale counts used in snake systematics heritable? SSAR/HL Annual Meeting. Raleigh, 
North Carolina. [Abstr]. 



Vol. 9, p. 160 Asiatic Herpetological Research 2001 

Thesis or dissertation 

Moody, S. 1980. Phylogenetic and historical biogeographical relationships of the genera in the Agamidae (Rep- 
tilia: Lacertilia). Ph.D. Thesis. University of Michigan. 373 pp. 

Anonymous, undated 

Anonymous. Undated. Turpan brochure. Promotion Department of the National Tourism Administration of the 
People's Republic of China, China Travel and Tourism Press, Turpan, Xinjiang Uygur Autonomous Region, 
China. 

Copyright 

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must be accompanied by the written consent of the copyright holder. 

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ures will be reduced to either 1 column (3.25") or two columns (6.5"). 

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NOTE: AHR now requires all tables to be submitted camera-ready. Please send 
hard copy, or electronic images of tables. Please use a sanserif font (Helvetica 
10pt or Ariel 10pt). 






Book Review: Wild About Reptiles. Field Guide to the Reptiles and Amphibians 

of the UAE 153 

Obituary: Sherman Anthony Minton Jr 154 

Guidelines for Manuscript Preparation and Submission 156 



Colophon. Asiatic Herpetological Research is created using Adobe FrameMaker 6, Acrobat 4, 
and Deneba Canvas 7 on Apple Macintosh computers. The body text is set in Times Roman 
and the headings in Helvetica. Using digital technology, we consumed less than 200 sheets of 
paper in the prepress production of this issue. 



ISSN 1051-382^ 



Wen-Jian Li. The Reproductive Bioiogy of Rana boulengeri 



Mingtao Song. Xiaomao Zeng, Guanfu Wu. Zhijun Liu, and Jinzhong Fu. A New Species of 
Batrachuperus from Northwestern China t> 

Rafe M. Brown, Alan E. Leviton, John W. Ferner. and Rogelio V. Sison. A New Snake of the 

Genus Hologerrhum Gunther (Reptilia; Squamata; Coluhridae) from Panay Island, Philippines 

9 

M. Farid Ahsan and Shayla Parvin. The First Record of Ptyas korros (Coluhridae) from 
Bangladesh 23 

Sean J. Blamires. Influence of Temperature on Burrow Use hy the Monitor Lizard Varanus 
panoptes of the Coastal Dunes at Fog Bay, Northern Australia 25 

Peter L. Cunningham. Notes on the Diet, Survival Rate, and Burrow Specifics of Uromastyx 
aegyptius microlepis from the United Arab Emirates 30 

John W. Ferner. Rafe M. Brown, Rogelio V. Sison, and Robert S. Kennedy. The Amphibians 
and Reptiles of Panay Island. Philippines 34 

James Ford Parham and Haitao Shi. The Discovery of Mauremys iversoni-like Turtles at a 
Turtle Farm in Hainan Province, China: The Counterfeit Golden Coin 71 

Matthias Stock, Daniel Frynta. Wolf-Riidiger Grosse, Claus Steinlein, and Michael Schmid. A 
Review of the Distribution of Diploid, Triploid and Tetraploid Green Toads (Bujo viridis 
complex) in Asia Including New Data from Iran and Pakistan 77 

Samraat Pawar and Sayantan Biswas. First Record of the Smooth-Backed Parachute Gecko 
Ptychozoon lionotum Annandale 1905 from the Indian Mainland 101 

Nasrullah Rastegar-Pouyam and Eskandar Rastegar-Pouyani. A New Species of Eremias 
(Sauria: Lacertidae) from Highlands of Kermanshah Province, Western Iran 107 

Konstantin A. Rogovin, Dmitry V. Semenov, and Georgy I. Shenbrot. Lizards of the Northern 
Mongolian Deserts: Densities and Community Structure 113 

Murat Sevinc and Ismail Hakki Ugurta§. The Morphology and Size of Blood Cells of Lacerta 
rudis bithynica 122 

Muhammad S. Khan. Buccopharyngeal Morphology and Feeding Ecology of Microhyla 
ornata tadpoles 1 30 

Ismail H. Ugurtas. Variation in Pelobates syriacus of Turkey 139 

Translation: Mingtao Song. A New Species of the Turtle Genus Cuora (Testudoformes: 
Testudinidae j 142 

Translation: Heinz Weissingcr. Testudo graeca anamurensis ssp. nov. from Asia Minor 145 

Book Review A Guide to th~ Fauna Of Iran . . 149 

Book Review: Four Receni Handbooks for Turkey .'...'. 1^1 



(Continued on inside of back cover) 



Harvard MCZ Llbran 



llll 

3 2044 066 300 427